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diff --git a/23354.txt b/23354.txt new file mode 100644 index 0000000..95355b9 --- /dev/null +++ b/23354.txt @@ -0,0 +1,22090 @@ +The Project Gutenberg eBook, Vegetable Teratology, by Maxwell T. Masters, +Illustrated by E. M. Williams + + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + + + + +Title: Vegetable Teratology + An Account of the Principal Deviations from the Usual Construction of Plants + + +Author: Maxwell T. Masters + + + +Release Date: November 6, 2007 [eBook #23354] + +Language: English + +Character set encoding: ISO-646-US (US-ASCII) + + +***START OF THE PROJECT GUTENBERG EBOOK VEGETABLE TERATOLOGY*** + + +E-text prepared by Charlene Taylor, Leonard Johnson, and the Project +Gutenberg Online Distributed Proofreading Team (https://www.pgdp.net) + + + +Note: Project Gutenberg also has an HTML version of this + file which includes the original illustrations. + See 23354-h.htm or 23354-h.zip: + (https://www.gutenberg.org/dirs/2/3/3/5/23354/23354-h/23354-h.htm) + or + (https://www.gutenberg.org/dirs/2/3/3/5/23354/23354-h.zip) + + +Transcriber's note: + + Words enclosed between equal signs were in bold face (=bold=). + + A list of corrections is at the end of the book. + + + + + +The Ray Society. +Instituted MDCCCXLIV. + +[Illustration] + +_This volume is issued to the Subscribers to the_ RAY SOCIETY _for +the Year_ 1868. + +London: +MDCCCLXIX. + +VEGETABLE TERATOLOGY, + +An Account of the Principal Deviations from the Usual +Construction of Plants + +by + +MAXWELL T. MASTERS, M.D., F.L.S. + +With Numerous Illustrations by E. M. Williams. + + + + + + + +London: +Published for the Ray Society by +Robert Hardwicke, 192. Piccadilly. +MDCCCLXIX. + + + + +TO +JOSEPH DALTON HOOKER, M.D., +D.C.L., LL.D., F.R.S., F.L.S., ETC. ETC. +DIRECTOR OF THE ROYAL GARDENS, KEW, + + +This Volume +IS +GRATEFULLY INSCRIBED. + + + + +TABLE OF CONTENTS. + + + PAGE + +INTRODUCTION xxi + + +BOOK I. + +DEVIATIONS FROM ORDINARY ARRANGEMENT 1 + + +PART I.--UNION OF ORGANS 8 + + +CHAPTER I. + +COHESION 9 + +Cohesion between axes of same plant, 9--Fasciation, 11--Cohesion +of foliar organs, 21--Of margins of single organs, 21--Tubular +petals, 23--Cohesion of several organs by their margins, 25--Of +the sepals, 27--Of the petals, 28--Of the stamens, 29--Of the +pistils, 29--Ascidia or pitchers, 30. + + +CHAPTER II. + +ADHESION 32 + +Adhesion of foliar organs, 32--Of leaves by their surfaces, +33--Of foliar to axile organs, 34--Of sepals to petals, 34--Of +stamens to petals, 34--Of stamens to pistils, 35--Miscellaneous +adhesions, 35--Of fruit to branch, 36--Synanthy, 36--Syncarpy, +45--Synspermy, 50--Between axes of different plants of same +species, 50--And of different species, 55--Synophty. + + +PART II.--INDEPENDENCE OF ORGANS 58 + + +CHAPTER I. + +FISSION 59 + +Fission of axile organs, 60--Of foliar organs, 61--Of petals, +66--Of stamens, 68--Of carpels, 68. + + +CHAPTER II. + +DIALYSIS 69 + +Dialysis of margins of individual parts, 70--Of margins of parts +of same whorl-calyx, 70--Of corolla, 71--Of stamens, 73--Of +carpels, 73. + + +CHAPTER III. + +SOLUTION 76 + +Solution of calyx from ovary, 77--Of stamens from petals, 82. + + +PART III.--ALTERATIONS OF POSITION 83 + + +CHAPTER I. + +DISPLACEMENT 84 + +Displacement of bulbs, 84--Of inflorescence, 85--Of leaves, +87--Of parts of flowers, 91--Of carpels, 96--Of placentas and +ovules, 96. + + +CHAPTER II. + +PROLIFICATION 100 + +Prolification of the inflorescence, 102--Median foliar, +103--Median floral, 105--Lateral foliar, 106--Lateral floral, +107--Prolification of the flower, 115--Median foliar, +116--Median floral, 119--Axillary prolification, 138--Foliar, +141--Floral, 142--Complicated prolification, 151--Of embryo, +155. + + +CHAPTER III. + +HETEROTAXY 156 + +Formation of adventitious roots, 156--Of shoots below the +cotyledons, 161--Adventitious leaves, 162--On scapes, +163--Production of leaves or scales in place of flower-buds, +164--Viviparous plants, 168--Formation of buds on leaves, +170--In pith, 171--On bulbs, 172--Production of gemmae in place +of spores, 173--Of flowers on leaves, 174--Of flower-buds in +place of leaf-buds, 176--Of flowers on spines, 177--Of +flower-buds on petals, 177--On fruits, 178--In ovaries, 180--Of +stamens in ovaries, 182--Of pollen in ovules, 185--Homomorphic +flowers of Compositae, 188--Heterotaxy affecting the +inflorescence, 188--Supra-soriferous ferns, 190. + + +CHAPTER IV. + +HETEROGAMY 190 + +Change in the position of male and female flowers, 191--From +monoecious to dioecious condition, 193--From dioecious to +monoecious, 193--From hermaphroditism to unisexuality, +195--From unisexuality to hermaphroditism, 197--Pollen replaced +by ovules, 201. + + +CHAPTER V. + +ALTERATIONS IN THE DIRECTION OF ORGANS 201 + +Fastigiation, 202--Eversion, 204--Altered direction of leaves, +205--Altered direction of flower, 206--Reflexion, +209--Gymnaxony, 211. + + +BOOK II. + +DEVIATIONS FROM ORDINARY FORM 213 + + +PART I.--STASIMORPHY 216 + + +CHAPTER I. + +PERSISTENCE OF JUVENILE FORMS 217 + +Stasimorphy in leaves of Conifers, 217--Regular peloria, 219. + + +PART II.--PLEIOMORPHY 228 + + +CHAPTER I. + +IRREGULAR PELORIA 228 + + +PART III.--METAMORPHY 240 + + +CHAPTER I. + +PHYLLODY 241 + +Phyllody of bracts, 242--In inflorescence of Conifers, 245--Of +calyx, 245--Of corolla, 251--Of stamens, 253--Of pistils, +256--Of ovules, 262--Changes in nucleus of ovule, 269--Phyllody +of accessory organs, 272--Chloranthy, 273--General remarks on, +278. + + +CHAPTER II. + +METAMORPHY OF FLORAL ORGANS 281 + +Sepalody of petals, 282--Petalody of calyx, calycanthemy, +283--Petalody of stamens, 285--Of anther, 291--Of connective, +293--Compound stamens, 294--Petalody of pistils, 296--Of ovules, +297--Of accessory organs, 297--Staminody of the bracts, 298--Of +sepals and petals, 298--Of pistils, 299--Of accessory organs, +301--Pistillody of perianth, 302--Of sepals, 302--Of stamens, +303--Of ovules, 310. + + +PART IV.--HETEROMORPHY 311 + + +CHAPTER I. + +DEFORMITIES 311 + +Formation of tubes, 312--In flower, 314--Spurs, 315--Contortion, +316--Spiral torsion, 319--Of leaf, 326--Adventitious tendrils, +326--Interrupted growth, 327--Cornute leaves, 328--Flattening, +328. + + +CHAPTER II. + +POLYMORPHY 329 + +Heterophylly, 330--Dimorphism, 333--Sports or bud-variations, +336. + + +CHAPTER III. + +ALTERATIONS OF COLOUR 337 + +Albinism, 337--Virescence, 338--Chromatism, 339. + + +BOOK III. + + +DEVIATIONS FROM ORDINARY NUMBER 340 + + +PART I.--INCREASED NUMBER OF ORGANS 343 + + +CHAPTER I. + +MULTIPLICATION OF AXILE ORGANS--INFLORESCENCE 346 + +Multiplication of branches, 346--Plica, 346--Polyclady, +347--Multiplication of branches of inflorescence, 348--Of bulbs, +350--Of florets, 351. + + +CHAPTER II. + +MULTIPLICATION OF FOLIAR ORGANS 352 + +Pleiophylly, 353--Multiplication of stipules, bracts, &c., +357--Polyphylly, 358--Increased number of leaves in a whorl, +358--Polyphylly of bracts, 358--Of calyx, 358--Of corolla, +359--Of androecium, 361--Of gynoecium, 363--Of flower in +general, 365--Increased number of ovules and seeds, 367--Of +embryos, 369--Of cotyledons, 370--Pleiotaxy, or multiplication +of whorls, 371--Pleiotaxy of bracts, 371--Of calyx, 374--Of +perianth, 375--Of corolla (hose in hose), 376--Androecium, +379--Androecium of Orchids, 380--Pleiotaxy of gynoecium, +388--Increased number of flowers in an inflorescence, 391. + + +PART II.--DIMINISHED NUMBER OF ORGANS 392 + + +CHAPTER I. + +SUPPRESSION OF AXILE ORGANS 393 + +Acaulosia, 393--Non-development of peduncle, 393--Nature of +calyx-tube (Casimir de Candolle), 394--Suppression of columella, +395. + + +CHAPTER II. + +SUPPRESSION OF FOLIAR ORGANS 395 + +Aphylly, 395--Meiophylly, 396--Of calyx or perianth, 396--Of +corolla, 397--Of androecium, 398--Of gynoecium, 399--Of +flower, 400--Meiotaxy, 403--Of calyx, 403--Of corolla, 403--Of +androecium, 405--Of gynoecium, 406--Of ovules or seeds, +407--Of flower, 408--General remarks on suppression, 409. + + +BOOK IV. + +DEVIATIONS FROM ORDINARY SIZE AND CONSISTENCE 413 + + +PART I--HYPERTROPHY 416 + + +CHAPTER I. + +ENLARGEMENT 417 + +Of axile organs, 418--Knaurs, 419--Enlargement of buds, 420--Of +flower-stalk, 421--In pears, 423--Of placenta, 424--Of leaves, +426--Development of parts usually abortive, 427--Enlargement of +perianth, 428--Of androecium, 430--Of gynoecium, 430--Of +fruit, 431--Alterations of consistence, 432. + + +CHAPTER II. + +ELONGATION 488 + +Elongation of root, 434--Of inflorescence, 434--Of +flower-stalks, 435--Of leaves, 437--Of parts of flower, 438--Of +thalamus and placenta, 440--Apostasis, 440. + + +CHAPTER III. + +ENATION 443 + +Excrescences from axile organs, warts, 444--Enation from leaves, +445--From sepals, 448--From petals, 448--Catacorolla, +451--Enation from stamens, 453--From carpels, 453. + + +PART II.--ATROPHY 454 + + +CHAPTER I. + +ABORTION 455 + +Abortion of axile organs, 455--Of receptacle, 457--Of leaves, +458--Of perianth, calyx, and corolla, 460--Of stamens, 463--Of +pollen, 463--Of pistil, fruit, &c., 464--Of ovules, +466--Depauperated ferns, 466--General remarks, 467. + + +CHAPTER II. + +DEGENERATION 470 + +Formation of scales, 470--Of hairs, 472--Of glands, 473--Of +tendrils, 473. + + +GENERAL CONCLUSIONS. + +General morphology of the leaf and axis, 476.--Homology, +476--Special morphology, 479--Calyx-tube, 480--Androecium, +482--Inferior ovary, 482--Placentation, 483--Structure of the +ovule, 484--Leaves of Conifers, 484--Relative position of +organs, 484--Law of alternation, 485--Co-relation, +486--Compensation, 488--Teratology and classification, 488. + + +APPENDIX. + +Double flowers, varieties of, 491--Causes of production, +491--Relation to variegated foliage, 497--List of plants +producing, 499. + + +NOTE 508 + +INDEX TO SUBJECTS 511 + +INDEX OF NAMES OF PLANTS 517 + + + + +LIST OF ILLUSTRATIONS. + + +FIG. PAGE + + 1. Diagram of regular pentamerous flower 4 + 2. Cohesion of branch _Dipsacus sylvestris_ 10 + 3. Fasciated lettuce ('Gard. Chron.') 11 + 4. Fasciation in _Asparagus_ 12 + 5. Fasciation in _Pinus Pinaster_ 13 + 6. Fasciation and spiral torsion in _Asparagus_ + ('Gard. Chron.') 14 + 7. Fasciation in scape of dandelion 16 + 8. Pitcher on leaf of _Pelargonium_ 22 + 9. Transition from flat to tubular segments of the perianth in + _Eranthis_ 24 + 10. Pitcher of _Crassula arborescens_ (C. Morren) 26 + 11. Gamopetalous corolla, _Papaver bracteatum_ 28 + 12. Adhesion of petaloid stamen to segment of perianth, + _Crocus_ 35 + 13. Adhesion of petal, stamen and carpel, + _Cheiranthus Cheiri_ 36 + 14. Synanthy. _Campanula Medium_ ('Gard. Chron.') 37 + 15, 16. Synanthic flowers of _Calanthe vestita_ 39 + 17. Synanthy in _Digitalis purpurea_ ('Gard. Chron.') 40 + 18. Synanthy in _Calceolaria_ 41 + 19, 20. Syncarpic apples 47 + 21. Adhesion of two stems of oak ('Gard. Chron.') 51 + 22. Adhesion of branches of elm ('Gard. Chron.') 52 + 23. Adhesion of two roots of carrot, one white, the other red + ('Gard. Chron.') 53 + 24. Section through inverted and adherent mushrooms 54 + 25. Bifurcated male catkin, _Cedrus Libani_ 61 + 26. Bifurcated leaf, _Lamium album_ 62 + 27. Bifurcated leaf, _Pelargonium_ 63 + 28. Bifurcated frond, _Scolopendrium_ 64 + 29. Three-lobed lip of _Oncidium_ 68 + 30. Dialysis of corolla in _Correa_ 71 + 31. Dialysis of corolla in _Campanula_ (De Candolle) 72 + 32. Anomalous form of orange 74 + 33, 34. Disjoined carpels of orange (Maout) 75 + 35. Proliferous rose (Bell Salter) 78 + 36, 37. Apple flower, with detached calyx, &c. 79 + 38. Flower of _OEnanthe crocata_, with detached calyx, &c. 80 + 39. Anomalous bulbs of tulip 85 + 40. Displaced leaf of _Gesnera_ (C. Morren) 88 + 41. Leaves of _Pinus pinea_ 89 + 42. Deranged leaves of yew 90 + 43. Cohesion of sepals and displacement of parts of _Oncidium + cucullatum_ 92 + 44. Malformed flower of _Cypripedium_ 93 + 45. Diagram of the same 93 + 46. Diagram of natural structure in _Cypripedium_ 93 + 47. Diagram of malformed flower of _Lycaste Skinneri_ 93 + 48. Diagram of malformed flower of _Dendrobium nobile_ 94 + 49. Natural arrangement in same flower 94 + 50. Diagram of malformed violet 94 + 51. Monstrous flower of _Cerastium_ ('Gard. Chron.') 97 + 52. Inflorescence of _Polyanthus_, with tufts of leaves + at the summit 105 + 53. Lateral prolification in inflorescence of _Pelargonium_ 108 + 54. Paniculate inflorescence of _Plantago major_ 109 + 55. Branched spike and leafy bracts of the same 110 + 56. Inflorescence of _Plantago lanceolata_, with leaves + and secondary flower-stalks at its summit 111 + 57. Branched inflorescence of _Reseda Luteola_ 112 + 58. Thalamus of strawberry prolonged into a leafy branch + ('American Agriculturist') 116 + 59. Flower of _Verbascum_, with dialysis of calyx and + corolla and prolonged thalamus 116 + 60. Median floral prolification of _Dianthus_ 120 + 61. Leafy carpels and prolification of _Daucus Carota_ 123 + 62. Median floral prolification of _Delphinium_ 126 + 63. Median prolification, &c., of _Orchis pyramidalis_ 128 + 64. Proliferous rose (Bell Salter) 130 + 65. Axillary floral prolification of _Nymphaea Lotus_ + ('Gard. Chron.') 144 + 66. Axillary floral prolification of _Dianthus_ 146 + 67. Proliferous rose 151 + 68. Proliferous rose ('Gard. Chron.') 152 + 69. Diagram of prolified orchid 153 + 70. Diagram of prolified orchis 154 + 71. Adventitious roots from petiole of celery 158 + 72. Germinating plant of mango 159 + 73, 74. Adventitious roots from leaves 160 + 75. Hip of rose bearing leaf 162 + 76. Leaves proceeding from the ovary of _Nymphaea_ sp. 162 + 77. Flower-stalk of dandelion, with leaves 163 + 78. Tuft of leaves in place of flowers in _Valeriana_ sp. 165 + 79. Scale-bearing spikelets of _Willdenovia_ 167 + 80. "Rose Willow" 167 + 81. Viviparous flowers of _Aira vivipara_ 169 + 82. Formation of shoot on leaf of _Episcia bicolor_ 171 + 83. Adventitious buds on root of sea-kale 172 + 84, 85. Production of adventitious bulbs in hyacinth ('Gard. + Chron.') 172 + 86. Adventitious buds on hyacinth ('Gard. Chron ') 173 + 87-92. Nepaul barley 174, 175 + 93. Formation of buds on fruit of _Opuntia_ 179 + 94. Flower-bud in the pod of _Sinapis_ 181 + 94*, 95. Adventitious pod in silique of _Cheiranthus_ 182 + 96, 97. Grapes, with adventitious fruits in interior 183 + 98. Stamens in ovary of _Baeckea diosmifolia_ 184 + 99. Pollen in ovule of passion flower (S. J. Salter) 185 +100. Female flowers at the summit of the inflorescence of + _Carex acuta_ 192 +101. Monoecious hop ('Gard. Chron.') 193 +102. Superior ovary, &c, of fuchsia 198 +103. Hermaphrodite flower of _Carica_ 199 +104. Ovuliferous anthers of _Cucurbita_ sp. 200 +105, 106. Prolonged inflorescence of fig 205 +107. Hollow turnip, with inverted leaves 206 +108. Normal flower of _Gloxinia_ 207 +109. Regular peloria of _Gloxinia_ 207 +110, 111. Structural details of erect _Gloxinia_ 208 +112. Reflected corolla of azalea 209 +113, 114. Enlarged and erect placenta of _Cuphea miniata_ + (C. Morren) 210 +115. Dimorphic leaves of _Juniperus sinensis_ 217 +116. Regular peloria of _Delphinium_ 219 +117. Structural details of the preceding 219 +118. Regular peloria of violet 220 +119. Double-flowered regular violet 220 +120. Regular peloria of _Eccremocarpus scaber_ 222 +121. Regular peloria of _Cattleya_ 223 +122. Peloria of _Calceolaria_ 230 +123. Peloric flower of _Aristolochia_ 232 +124, 125. Peloric flowers of _Corydalis_ 236 +126. Rose plantain 242 +127. Leafy bracts in _Plantago major_ 243 +128. Leafy scales of _Dahlia_ 244 +129. Leafy sepals of rose 246 +130. Leafy sepals of Fuchsia ('Gard. Chron.') 247 +131. Leafy calyx of primrose 248 +132. Leafy calyx of melon 248 +133. Leafy sepals and petals of _Geranium_ 251 +134. Leafy stamens of _Petunia_ 254 +135, 136. Leafy anthers of _Jatropha_ (Mueller) 255 +137. Proliferous rose, with leafy carpels, &c. (Bell Salter) 257 +138. Cucumber, with adventitious leaf attached (S. J. Salter) 258 +139. Leafy carpels in flower of _Triumfetta_ 260 +140. Leafy ovules of _Sinapis_ 264 +141, 142. Leafy ovules of _Trifolium repens_ (Caspary) 265 +143. Portion of leafy carpel of _Delphinium_, with ovules + (Cramer) 266 +144, 145. Enlarged view of section of leafy carpel, &c., of + _Delphinium_ (Cramer) 267 +146. Placentae of _Dianthus_, bearing ovules and carpels + ('Gard. Chron.') 268 +147. Ovules passing into carpels, _Dianthus_ + ('Gard. Chron.') 268 +148. Leafy shoot in place of ovule of _Gaillardia_ 270 +149, 150. Leafy sepals, petals, &c., of _Epilobium_ 273 +151. Leafy carpel of rose, with deformed ovules 274 +152. Flower of St. Valery apple 282 +153. Petaloid calyx of _Mimulus_ 284 +154. Double stellate columbine 287 +155. Four-winged filaments of Rhododendron 290 +156. Diagram of malformed flower of _Catasetum_ 291 +157. Petaloid stamen of _Viola_ 292 +158. Double columbine 293 +159. Petaloid stamens of _Hibiscus_ 293 +160. Displaced coloured leaf, &c., of tulip 302 +161. Supernumerary carpels in orange (Maout) 303 +162. Pistilloid stamens of poppy 304 +163. Pistilloid stamens of wallflower, &c. 306 +164. Passage of stamen to carpel in lily 307 +165. Transition of stamens to carpels, + _Sempervivum tectorum_ 309 +166. Ascidia of cabbage 312 +167. Stalked pitcher on lettuce leaf 313 +168. Tubular petal of _Primula sinensis_ 315 +169. Spurs on flower of _Calceolaria_ 316 +170. Contorted stem of _Juncus_ 317 +171. Contorted branch of _Crataegus_ 317 +172. Spirally-twisted stem of teazel 321 +173. Spirally-twisted stem _Galium_ 323 +174. Spirally-twisted root ('Gard. Chron.') 324 +175. Interrupted growth of radish (American Agriculturist) 327 +176. Interrupted growth in apple 327 +177. Polymorphous leaves of lilac 331 +178. Adventitious growth on frond of _Pteris quadriaurita_ 333 +179, 180. Coloured flower-stalks of feather hyacinth + (C. Morren) 347, 348 +181. Multiplication of catkins, _Corylus_ 349 +182. Branched inflorescence of broccoli ('Gard. Chron.') 351 +183, 184. Supernumerary leaf of elm 353, 354 +185. Supernumerary leaf of hazel 355 +186. Multiplication of parts of flower in a plum ('Gard. Chron.') 366 +187. Wheat-ear carnation 372 +188. Multiplication of bracts in _Delphinium Consolida_ 373 +189. Multiplication of bracts in _Pelargonium_ 373 +190. Double white lily 376 +191. Double flower of _Campanula rotundifolia_ 378 +192. Diagram of usual arrangement of parts in _Orchis_ + (Darwin) 381 +193. Diagram of malformed flower of _Ophrys aranifera_ 385 +194. Malformed flower of _Ophrys aranifera_ 385 +195. Diagram of malformed flower of _Orchis mascula_ + (Cramer) 386 +196. Multiplication of carpels, Tulip 388 +197. Section of St. Valery apple 388 +198. Regular dimerous flower of _Calanthe vestita_ 402 +199. Regular dimerous flower of _Odontoglossum Alexandrae_ 402 +200. Hypertrophied branch of _Pelargonium_ 418 +201. Tubers in the axils of leaves of the potato 420 +202. Hypertrophied pedicels of ash 421 +203, 204. Hypertrophy and elongation of flower-stalk, &c., in + pears 422, 423 +205. Hypertrophied perianth, _Cocos nucifera_ 428 +206. Elongation of flower-stalk, _Ranunculus acris_ 436 +207. Linear leaf-lobes of parsley 438 +208. Passage of pinnate to palmate leaves in horse-chestnut 439 +209. Elongation of thalamus, apostasis, &c., in flower of + _Delphinium_ (Cramer) 441 +210. Adventitious growths from cabbage leaf 445 +211. Crested fronds of _Nephrodium molle_ 447 +212. Supernumerary petals, &c., _Datura fastuosa_ 450 +213. Supernumerary petaloid segments in flower of _Gloxinia_ 451 +214. Catacorolla of _Gloxinia_ (E. Morren) 452 +215. Atrophied leaves of cabbage 460 +216. Abortion of petals, pansy 461 +217. Flower of _Oncidium abortivum_ 462 +218. Bladder plum 464 + + + + +INTRODUCTION. + + +Till within a comparatively recent period but little study was given to +exceptional formations. They were considered as monsters to be shunned, +as lawless deviations from the ordinary rule, unworthy the attention of +botanists, or at best as objects of mere curiosity. By those whose +notions of structure and conformation did not extend beyond the details +necessary to distinguish one species from another, or to describe the +salient features of a plant in technical language; whose acquaintance +with botanical science might almost be said to consist in the +conventional application of a number of arbitrary terms, or in the +recollection of a number of names, teratology was regarded as a chaos +whose meaningless confusion it were vain to attempt to render +intelligible,--as a barren field not worth the labour of tillage. + +The older botanists, it is true, often made them the basis of satirical +allusions to the political or religious questions of the day, especially +about the time of the Reformation, and the artists drew largely upon +their polemical sympathies in their representations of these anomalies. +Linnaeus treated of them to some extent in his 'Philosophia,' but it is +mainly to Angustin Pyramus De Candolle that the credit is due of calling +attention to the importance of vegetable teratology. This great +botanist, not only indirectly, but from his personal research into the +nature of monstrosities, did more than any of his predecessors to rescue +them from the utter disregard, or at best the contemptuous indifference, +of the majority of botanists. De Candolle gave a special impetus to +morphology in general by giving in his adhesion to the morphological +hypotheses of Goethe. These were no mere figments of the poet's +imagination, as they were to a large extent based on the actual +investigation of normal and abnormal organisation by Goethe both alone, +and also in conjunction with Batsch and Jaeger. + +De Candolle's example was contagious. Scarcely a botanist of any +eminence since his time but has contributed his quota to the records of +vegetable teratology, in proof of which the names of Humboldt, Robert +Brown, the De Jussieus, the Saint Hilaires, of Moquin-Tandon, of +Lindley, and many others, not to mention botanists still living, may be +cited. To students and amateurs the subject seems always to have +presented special attractions, probably from the singularity of the +appearances presented, and from the fact that in many cases the +examination of individual instances of malformation can be carried on, +to a large extent, without the lengthened or continuous investigation +and critical comparative study required by other departments of +botanical science. Be this as it may, teratology owes a very large +number of its records to this class of observers. + +While the number of scattered papers on vegetable teratology in various +European languages is so great as to preclude the possibility of +collating them all, there is no general treatise on the subject in the +English language, with the exception of Hopkirk's 'Flora Anomala,' a +book now rarely met with, and withal very imperfect; and this +notwithstanding that Robert Brown early lent his sanction to the +doctrines of Goethe, and himself illustrated them by teratological +observations. In France, besides important papers of Turpin, Geoffroy de +Saint Hilaire, Brongniart, Kirschleger and others, to which frequent +allusion is made in the following pages, there is the classic work of +Moquin-Tandon, which was translated into German by Schauer. Germany has +also given us the monographs of Batsch, Jaeger, Roeper, Engelmann, +Schimper, Braun, Fleischer, Wigand, and many others. Switzerland has +furnished the treatises of the De Candolles, and of Cramer; Belgium, +those of Morren, &c., all of which, as well as many others that might be +mentioned, are, with the exception of Moquin-Tandon's 'Elements,' to be +considered as referring to limited portions only and not to the whole +subject.[1] + +In the compilation of the present volume great use has been made of the +facts recorded in the works just cited, and especially in those of +Moquin-Tandon, Engelmann, and Morren. A very large number of +communications on teratological subjects in the various European +scientific publications have also been laid under contribution. In most +cases reference has been given to, and due acknowledgment made of, the +sources whence information has been gathered. Should any such reference +be omitted, the neglect must be attributed to inadvertence, not to +design. In selecting illustrations from the immense number of recorded +facts, the principle followed has been to choose those which seemed +either intrinsically the most important, or those which are recorded +with the most care. In addition to these public sources of information, +the author has availed himself of every opportunity that has offered +itself of examining cases of unusual conformation in plants. For many +such opportunities the author has to thank his friends and +correspondents. Nor has he less reason to be grateful for the +suggestions that they have made, and the information they have supplied. +In particular the writer is desirous of acknowledging his obligations to +the Society, under whose auspices this work is published, and to Mr. S. +J. Salter, to whom the book in some degree owes its origin. + +The drawings, where not otherwise stated, have been executed either from +the author's own rough sketches, or from the actual specimens, by Mr. E. +M. Williams. A large number of woodcuts have also been kindly placed at +the disposal of the author by the proprietors of the 'Gardeners' +Chronicle.'[2] + + +As it is impossible to frame any but a purely arbitrary definition of +teratology or to trace the limits between variation and malformation, it +may suffice to say that vegetable teratology comprises the history of +the irregularities of growth and development in plants, and of the +causes producing them. These irregularities differ from variations +mainly in their wider deviation from the customary structure, in their +more frequent and more obvious dependence on external causes rather than +on inherent tendency, in their more sudden appearance, and lastly in +their smaller liability to be transmitted by inheritance. + +What may be termed normal morphology includes the study of the form, +arrangement, size and other characteristic attributes of the several +parts of plants, their internal structure, and the precise relation one +form bears to another. In order the more thoroughly to investigate these +matters it is necessary to consider the mode of growth, and specially +the plan of evolution or development of each organ. This is the more +needful owing to the common origin of things ultimately very different +one from the other, and to the presence of organs which, in the adult +state, are identical or nearly so in aspect, but which nevertheless are +very unlike in the early stages of their existence.[3] Following Goethe, +these changes in the course of development are sometimes called +metamorphoses. In this way Agardh[4] admits three kinds of +metamorphosis, which he characterises as: 1st. Successive metamorphoses, +or those changes in the course of evolution which each individual organ +undergoes in its passage from the embryonic to the adult condition, or +from the simple and incomplete to the complex and perfect. 2. Ascending +metamorphoses, including those changes of form manifested in the same +adult organism by the several parts of which it consists--those parts +being typically identical or homologous, such as the parts of the +flower, or, in animals, the vertebrae, &c. 3. Collateral metamorphoses, +comprising those permutations of form and function manifested in +homologous organs in the different groups of organisms, classes, orders, +genera, species, &c. + +Thus, in the first instance, we have a comparative examination of the +form of each or any separate part of the same individual at different +epochs in its life-history; in the second we have a similar comparison +instituted between the several parts of the same organism which +originally were identical in appearance, but which have in course of +evolution altered in character. In the third form we have the +comparative view not of one organ at different times, nor of the several +parts of one organism, but of the constituent elements pertaining to +those aggregates of individuals to which naturalists apply the terms +classes, orders, &c. + +In successive metamorphosis we have a measure of the amount of change +and of the perfection of structure to which each separate organ attains. + +In ascending metamorphosis we have a gauge of the extent of alteration +that may take place in the several homologous organs under existing +circumstances. + +In collateral metamorphosis, in the same way, we have an illustration of +the degree of change possible in aggregates of organisms under existing +circumstances. + +Now it is clear that from an investigation of all three classes just +mentioned, we shall be able to gain an idea of those points which are +common to all parts, to all individuals or to all aggregates, and those +that are peculiar to some of them, and, by eliminating the one from the +other, we shall arrive at conclusions which will be more or less +generally accurate or applicable, according to the ability of the +student and the extent to which the comparative analysis is earned. It +is thus that morphologists have been enabled to frame types or standards +of reference, and systematists to collocate the organisms they deal with +into groups. These standards and groups are more or less artificial +(none can be entirely natural) in proportion to the amount of knowledge +possessed by their framers, and the use they make of it. + +From this point of view teratological metamorphosis of all three kinds +demands as much attention as that which is called normal. We can have no +thorough knowledge of an organ, of an individual which is an aggregate +of organs, or of an aggregate of individuals of whatever degree, unless +we know approximately, at least, what are the limits of each. It is not +possible to trace these limits accurately in the case of natural +science, but the larger our knowledge and the wider our generalisations, +the closer will be our approach to the truth. + + +The most satisfactory classification of malformations would be one +founded upon the nature of the causes inducing the several changes. +Thus, in all organised beings, there is a process of growth, mere +increase in bulk as it were, and a process of evolution or +metamorphosis, in accordance with which certain parts assume a +different form from the rest, in order the better to fit them for the +performance of different offices. Should growth and development be +uniform and regular, that is in accordance with what is habitual in any +particular species, there is no monstrosity, but if either growth or +development be in any way irregular, malformation results. Hence, +theoretically, the best way of grouping cases of malformation would be +according as they are the consequences of:--1st. Arrest of Growth; +2ndly. Excessive Growth; 3rdly. Arrest of Development; 4thly, of +Excessive or Irregular Development. + +In practice, however, there are so many objections to this plan that it +has not been found practicable to carry it out. The inability arises to +a great extent from our ignorance of what should be attributed to arrest +of growth, what to excess of development, and so on. Moreover, a student +with a malformed plant before him must necessarily ascertain in what way +it is malformed before he can understand how it became so, and for this +purpose any scheme that will enable him readily to detect the kind of +monstrosity he is examining, even though it be confessedly artificial +and imperfect will be better than a more philosophical arrangement which +circumstances prevent him from employing. + +The plan followed in this volume is a slight modification of that +adopted by Moquin-Tandon, and with several additions. In it the aim is +to place before the student certain salient and easily recognisable +points by reference to which the desired information can readily be +found. Under each subdivision will be found general explanatory remarks, +illustrative details, and usually a summary of the more important facts +and the inferences to be derived from them. Bibliographical references +and lists of the plants most frequently affected with particular +malformations are also given. In reference to both these points it must +be remembered that absolute completeness is not aimed at; had such +fullness of detail been possible of attainment it would have +necessitated for its publication a much larger volume than the +present.[5] It is hoped that both the lists of books and of plants are +sufficiently full for all general purposes.[6] + +In the enumeration of plants affected with various malformations the ! +denotes that the writer has himself seen examples of the deviation in +question in the particular plant named, while the prefix of the * +indicates that the malformation occurs with special frequency in the +particular plant to which the sign is attached. + +Teratological alterations are rarely isolated phenomena, far more +generally they are associated with other and often compensatory changes. +Hence it is often necessary, in studying any given malformation, to +refer to two or more subdivisions, and in this way a certain amount of +repetition becomes unavoidable. The details of the several cases of +malformation given in these pages are generally arranged according to +their apparent degree of importance. Thus, in a case of prolification +associated with multiplication of the petals, the former change is a +greater deviation from the customary form than the latter, hence +reference should be made, in the first instance, to the sections +treating on prolification, and afterwards to those on multiplication. To +facilitate such research, numerous cross references are supplied. + +In the investigation of teratological phenomena constant reference must +be made to the normal condition, and _vice versa_, else neither the one +nor the other can be thoroughly understood. It cannot, however, be +overlooked that the form and arrangement called normal are often merely +those which are the most common, while the abnormal or unusual +arrangement is often more in consonance with that considered to be +typical than the ordinary one. Thus, too, it is often found that the +structural arrangements, which in one flower are normal, are in another +abnormal, in so far that they are not usual in that particular instance. + +For purposes of reference, a standard of comparison is required; and +this standard, so long as its nature is not overlooked, may, indeed must +be, to some extent, an arbitrary one. Thus in the phanerogamous plants +there is assumed to exist, in all cases, an axis (stem, branches, roots, +thalamus, &c.), bearing leaves and flowers. These latter consist of four +whorls, calyx, corolla, stamens, and pistils, each whorl consisting of +so many separate pieces in determinate position and numbers, and of +regular proportionate size. A very close approach to such a flower +occurs normally in _Limnanthes_ and _Crassula_, and, indeed, in a large +proportion of all flowers in an early stage of development. To a +standard type, such as just mentioned, all the varied forms that are met +with, either in normal or abnormal morphology, may be referred by +bearing in mind the different modifications and adaptations that the +organs have to undergo in the course of their development. Some parts +after a time may cease to grow, others may grow in an inordinate degree, +and so on; and thus, great as may be the ultimate divergences from the +assumed standard, they may all readily be explained by the operation, +simply or conjointly, of some of the four principal causes of +malformation before alluded to. The fact that so many and such varied +changes can thus readily be explained is not only a matter of +convenience, but may be taken as evidence that the standard of reference +is not wholly arbitrary and artificial, but that it is a close +approximation to the truth. + +It has already been said that an arrangement like that here considered +as typical is natural to some flowers in their adult state, and to a +vast number in their immature condition. It would be no extravagant +hypothesis to surmise that this was the primitive structure of the +flower in the higher plants. Variations from it may have arisen in +course of time, owing to the action of an inherent tendency to vary, or +from external circumstances and varied requirements which may have +induced corresponding adaptations, and which may have been transmitted +in accordance with the principle of hereditary transmission. This +hypothesis necessarily implies a prior simplicity of organisation, of +which, indeed, there is sufficient proof; many cases of malformation +can thus be considered as so many reversions to the ancestral form. + +Thus, teratology often serves as an aid in the study of morphology in +general, and also in that of special groups of plants, and hence may +even be of assistance in the determination of affinities. In any case +the data supplied by teratology require to be used with caution and in +conjunction with those derived from the study of development and from +analogy. It is even possible that some malformations, especially when +they acquire a permanent nature and become capable of reproducing +themselves by seed, may be the starting-point of new species, as they +assuredly are of new races, and between a race and a species he would be +a bold man who would undertake to draw a hard and fast line.[7] + +Discredit has been cast on teratology because it has been incautiously +used. At one time it was made to prove almost everything; what wonder +that by some, now-a-days, it is held to prove nothing. True the evidence +it affords is sometimes negative, often conflicting, but it is so rather +from imperfect interpretation than from any intrinsic worthlessness. If +misused the fault lies with the disciple, not with Nature. + +Teratology as a guide to the solution of morphological problems has been +especially disparaged in contrast with organogeny, but unfairly so. +There is no reason to exalt or to disparage either at the expense of the +other. Both should receive the attention they demand. The study of +development shows the primitive condition and gradual evolution of parts +in any given individual or species; it carries us back some stages +further in the history of particular organisms, but so also does +teratology. Many cases of arrest of development show the mode of growth +and evolution more distinctly, and with much greater ease to the +observer, than does the investigation of the evolution of organs under +natural circumstances. Organogeny by no means necessarily, or always, +gives us an insight into the principles regulating the construction of +flowers in general. It gives us no archetype except in those +comparatively rare cases where primordial symmetry and regularity exist. +When an explanation of the irregularity of development in these early +stages of the plant's history is required, recourse must be had to the +inferences and deductions drawn from teratological investigations and +from the comparative study of allied forms precisely as in the case of +adult flowers. + +The study of development is of the highest importance in the examination +of plants as individuals, but in regard to comparative anatomy and +morphology, and specially in its relation to the study of vegetable +homology it has no superiority over teratology. Those who hold the +contrary opinion do so, apparently, because they overlook the fact that +there is no distinction, save of degree, to be drawn between the laws +regulating normal organisation, and those by which so-called abnormal +formations are regulated. + +It is sometimes said, and not wholly without truth, that teratology, as +it stands at present, is little more than a record of facts, but in +proportion as the laws that regulate normal growth are better +understood, so will the knowledge of those that govern the so-called +monstrous formations increase. Sufficient has been already said to +prove that there is no intrinsic difference between the laws of growth +in the two cases. As our knowledge increases we shall be enabled to +ascertain approximately of what extent of variation a given form is +capable, under given conditions, and to refer all formations now +considered anomalous to a few well-defined forms. Already teratology has +done much towards showing the erroneous nature of many morphological +statements that still pass current in our text-books, though their +fallacy has been demonstrated again and again. Thus organs are said to +be fused which were never separate, disjunctions and separations are +assigned to parts that were never joined, adhesions and cohesions are +spoken of in cases where, from the nature of things, neither adhesion +nor cohesion could have existed. Some organs are said to be atrophied +which were never larger and more fully developed than they now are, and +so on. So long as these expressions are used in a merely conventional +sense and for purposes of artificial classification or convenience, well +and good, but let us not delude ourselves that we are thus contributing +to the philosophical study either of the conformation of plants or of +the affinities existing between them. What hope is there that we shall +ever gain clear conceptions as to the former, as long as we tie +ourselves down to formulas which are the expressions of facts as they +appear to be, rather than as they really are? What chance is there of +our attaining to comprehensive and accurate views of the genealogy and +affinities of plants as long as we are restricted by false notions as to +the conformation and mutual relation of their parts?[8] + +That teratology may serve the purposes of systematic botany to a greater +extent than might at first be supposed becomes obvious from a +consideration of such facts as are mentioned under the head of Peloria, +while the presence of rudimentary organs, or the occasional appearance +of additional parts, or other changes, may, and often do, afford a clue +to the relationship existing between plants--a relationship that might +otherwise be unsuspected. So, too, some of the alterations met with +appear susceptible of no other explanations than that they are +reversions to some pre-existing form, or, at any rate, that they are +manifestations of a phase of the plant affected different from that +which is habitual, and due, as it were, to a sort of allotropism. + +The mutations and perversions of form, associated as they commonly are +with corresponding changes of function, show the connection between +teratology and physiology--a connection which is seen to be the more +intimate when viewed in the light afforded by the writings and +experiments of Gaertner, Sprengel, and St. Hilaire, and, in our own +times, especially by the writings and experiments of Mr. Darwin, whose +works on the 'Origin of Species,' and particularly on the 'Variation of +Animals and Plants under Domestication' comprise so large a collection +of facts for the use of students in most departments of biology. It +will suffice to allude, in support of these statements, to the writings +of Mr. Darwin on such subjects as rudimentary organs, the use or disuse +of certain parts according to circumstances, the frequently observed +tendency of some flowers to become structurally unisexual, the liability +of other flowers perfectly organised to become functionally imperfect, +at least so far as any reciprocal action of the organs of the same +flower is concerned, reversions, classification, general morphology, and +other subjects handled at once with such comprehensive breadth and +minute accuracy of detail by our great physiologist. + +In the following pages alterations of function, unless attended by +corresponding alterations of form, are either only incidentally alluded +to, or are wholly passed over; such, for instance, as alterations in the +period of flowering, in the duration of the several organs, and so +forth.[9] Pathological changes, lesions caused by insect puncture or +other causes, also find no place in this book, unless the changes are of +such a character as to admit of definite comparison with normal +conformation. Usually such changes are entirely heteromorphous, and, as +it were, foreign to the natural organisation. + +The practical applications of teratology deserve the attention of those +cultivators who are concerned in the embellishment of our gardens and +the supply of our tables. The florist lays down a certain arbitrary +standard of perfection, and attempts to make flowers conform to that +model. Whether it be in good taste or not to value all flowers, in +proportion as they accord with an artificial and comparatively inelastic +standard of this kind, we need not stop to enquire; suffice it to say, +that taking the matter in its broadest sense, the aim of the florist is +to produce large, symmetrical flowers, brightly and purely coloured, or +if parti-coloured, the colours must be distinct, harmonious, or +contrasted. When all this is done, the flower, in most instances, +becomes 'monstrous' of the eyes in the botanist, though all the more +interesting to the student of morphology on that account. In like manner +the double flowers, the "breaks," the "sports" which the florist +cultivates so anxiously, are all of them greater or less deviations from +the ordinary form, while the broccolies, the cabbages, and many other +products of our kitchen gardens and fields owe the estimation in which +they are held entirely to those peculiarities which, by an unhappy +application of words, are called monstrous by botanists. Grafting, +layering, the "striking" of cuttings, the formation of adventitious +roots and buds, processes on which the cultivator so greatly relies for +the propagation and extension of his plants, are also matters with which +teratology concerns itself. Again the difficulty experienced +occasionally in getting vines, strawberries, &c., to set properly, may +sometimes be accounted for by that inherent tendency which some plants +possess of exchanging an hermaphrodite for a unisexual condition. + +For reasons then of direct practical utility, no less than on purely +scientific grounds, it is desirable to study these irregularities of +growth, their nature, limits, and inducing causes; and to this end it is +hoped the present work may, in some degree, contribute. + +FOOTNOTES: + +[1] An excellent summary of the history of Vegetable Teratology is given +in Kirschleger's 'Essai historique de la Teratologie Vegetale,' +Strasburg, 1845. + +[2] In some instances diagrams and formulae are given in explanation of +the conformation of monstrous flowers; in general these require no +further explanation than is given in the text, unless it be to state +that the horizontal line--is intended to indicate the cohesion of the +parts over which it is placed, while the vertical line | signifies the +adhesion of the organs by whose side it is placed. The formula + + S S S S S + ------------------------ + | P P P P P + | + | ST ST ST ST ST + +shows that the sepals (S) are distinct, the petals (P) coherent, and the +stamens (ST) adherent to the petals. + +[3] Wolff was the first to call attention to the great importance of the +study of development. He was followed by Turpin, Mirbel, Schleiden, +Payer, and others, and its value is now fully recognised by botanists. + +[4] Agardh, "Theoria Syst. Plant.," p. xxiii. + +[5] In the memoirs of Hopkirk, Kirschleger, Cramer, Hallier, and others, +malformations are arranged primarily according to the organs affected, +an arrangement which has only convenience to justify it. It is hoped +that the index and the headings to the paragraphs in the present volume +will suit the convenience of the reader as well as if the more +artificial plan just alluded to had been adopted. + +[6] Cryptogamous plants are only incidentally alluded to in these pages, +owing to their wide difference in structure from flowering plants. +Attention may, also, here be called to a paper of M. de Seynes in a +recent number of the Bulletin of the Botanical Society of France, vol. +xiv, p. 290, tab. 5 et 6, in which numerous cases of malformation among +agarics are recorded. See also same publication, vol. iv, p. 744; vol. +v, p. 211; vol. vi, p. 496. + +[7] On this subject see a paper of M. Naudin in the 'Comptes Rendus,' +1867, t. 64, pp. 929-933. + +[8] It is probable that many terms and expressions calculated to mislead +in the way above mentioned are made use of in the following pages. The +inconsistency manifested by their use may be excused on the ground of +ignorance of the true structure, and by the circumstance that in many +cases facts alone are recorded without an explanation of them being +offered. Moreover, it is desirable to act in conformity with the usual +practice of botanical writers, and not to change established +terminology, even if suspected to convey false ideas, until the true +condition of affairs be thoroughly well ascertained by organogenetic +research or other means. + +[9] A curious illustration of the latter class of alterations came under +the writer's notice last summer (1868), and which he has reason to +believe has not been previously recorded, viz. the persistence in an +unwithered state of the petals at the base of the ripe fruit, in a +strawberry. All the fruits on the particular plants alluded to were thus +provided as it were with a white frill. Whether this be a constant +occurrence in the particular variety is not known. + + + + +VEGETABLE TERATOLOGY. + + + + +BOOK I. + +DEVIATIONS FROM THE ORDINARY ARRANGEMENT OF ORGANS. + + +As full details relating to the disposition or arrangement of the +general organs of flowering plants are given in all the ordinary +text-books, it is only necessary in this place to allude to the main +facts at present known, and which serve as the standard of comparison +with which all morphological changes are compared. + +Even in the case of the roots, which appear to be very irregular in +their ramification, it has been found that, in the first instance at +least, the rootlets or fibrils are arranged in regular order one over +another, in a certain determinate number of vertical ranks, generally +either in two or in four, sometimes in three or in five series. This +regularity of arrangement (Rhizotaxy), first carefully studied by M. +Clos, is connected with the disposition of the fibro-vascular bundles in +the body of the root. This primitive regularity is soon lost as the +plant grows. + +In the case of the leaves there are two principal modes of arrangement, +dependent, as it would seem, on their simultaneous or on their +successive development; thus, if two leaves on opposite sides of the +stem are developed at the same time, we have the arrangement called +opposite; if there are more than two, the disposition is then called +verticillate or whorled. On the other hand, if the leaves are developed +in succession, one after the other, they are found to emerge from the +stem in a spiral direction. In either case the leaves are arranged in a +certain regular manner, according to what are called the laws of +Phyllotaxis, which need not be entered into fully here; but in order the +better to estimate the teratological changes which take place, it may be +well to allude to the following circumstances relating to the +alternation of parts. The effect of this alternation is such, that no +two adjacent leaves stand directly over or in front one of the other, +but a little to one side or a little higher up. Now, in the alternate +arrangement the successive leaves of each spiral cycle alternate one +with another till the coil is completed. For the sake of clearness this +may be illustrated thus:--Suppose the spiral cycle to comprise five +leaves, numbered 1, 2, 3, 4, 5, then 2 would intervene between 1 and 3, +and so on, while the sixth leaf would be the commencement of a new +series, and would be placed exactly over 1. This arrangement may be thus +formularised: + + 6 7 8 9 10 + 1 2 3 4 5 + +In the verticillate or simultaneous arrangement of leaves the case is +somewhat different. Let us suppose a whorl of eight leaves, surmounted +by a similar whorl of eight. In such a case it will generally be found +that the whorls alternate one with another, as may be represented by +this symbol: + + 9 10 11 12 13 14 15 16 + 1 2 3 4 5 6 7 8 + +The simplest illustration of this arrangement is seen in the case of +decussate leaves, where those organs are placed in pairs, and the pairs +cross one another at right angles. This may be expressed by the +following symbol: + + 7 8 + 5 6 + 3 4 + 1 2 + +Thus, while in both the annular and the spiral modes of development the +individual members of each complete series necessarily alternate one +with another, in the former case the series themselves alternate, while +in the successive arrangement they are placed directly one over the +other. There are, of course, exceptions, but the rule is as has been +stated, and the effect is to prevent one leaf from interfering with the +development and growth of its neighbours. + +In the case of the whorled or simultaneous arrangement the conditions of +growth must be uniform on all sides, but in the successive or spiral +disposition the conditions influencing growth act with unequal force, on +different sides of the stem, at the same time. In the whorl there is an +illustration of radiating symmetry, while in the spiral arrangement +there is a transition to the bilateral symmetry. There are frequent +passages from one to the other even under normal circumstances; thus, +while the one arrangement obtains in the ordinary leaves, the parts of +the flower may be disposed according to the other method. In the annular +disposition it generally happens that the rings are separated one from +the other by the development of the stem between them, the internodes +between the constituent leaves themselves of course being undeveloped; +on the other hand, in the spiral or successive arrangement there is no +such alternate growth and arrest of growth of the stem between the +leaves, or between successive cycles, but the growth is, under favorable +conditions, continuous--leaf is separated from leaf, and cycle from +cycle, by the continually elongating stem. Thus, the two modes of growth +correspond precisely with those observed in the case of definite and +indefinite inflorescence respectively. + +[Illustration: FIG. 1.--Diagram showing the arrangement of parts in a +complete, regular, pentamerous flower: _s_, sepals; _p_, petals; _st_, +stamens; _o_, ovaries.] + +The same arrangements, that are observed in the disposition of the +leaves, apply equally well to the several parts of the flower; thus, in +what is for convenience considered the typical flower, there is a calyx +of five or more distinct sepals, equal in size, and arranged in a whorl, +a corolla of a similar number of petals alternating with the sepals, +five stamens placed in the same position with reference to the petals, +and five carpels alternating with the stamens. Throughout this book this +arrangement is taken as the standard of reference. Nevertheless the +spiral order does occur in the floral leaves as well as in those of the +stem; it often happens, especially when the organs are numerous, that +they form spiral series; and the same holds good very generally, when +the parts of the flower are uneven in number, as in the very common +quincuncial arrangement of the sepals, &c. + +To these general remarks, intended to show the agreement between the +disposition of the leaves of the stem and those of the flower, it is +merely necessary to add that the arrangement of the placentas, as well +as that of the ovules borne on them, is also definite, and takes place +according to methods explained in all the text-books, and on which, +therefore, it is not necessary to dilate in this place. + +The branches of the stem or axis correspond for the most part in +disposition with that of the leaves from the axils of which they +originate, subject, however, to numerous disturbing causes, and to +alterations from the usual or typical order brought about by the +development of buds. These latter organs, as it seems, may be found in +almost any situation, though their ordinary position is in the axil of a +leaf or at the end of a stem or branch. + +The points just mentioned are of primary importance in structural +botany, and as such are seized on not only by the morphologist, but by +the systematic botanist, who finds in them the characters by which he +may separate one group from another. Thanks to the labours of those +observers who have devoted their attention to that difficult but most +important branch of study, organogeny, or the investigation of the +development of the various organs, and to the researches of the students +of comparative anatomy or morphology, the main principles regulating the +arrangement and form of the organs of flowering plants seem to be fairly +well established, though in matters of detail much remains to be +cleared up, even in such important points as the share which the axis +takes in the construction of the flower and fruit, the nature of the +placenta, the construction of the ovules, and other points. + +The facts already known justify the adoption of a standard or typical +arrangement as just mentioned. The intrinsic value of this type is shown +by the facility with which all varieties of form or arrangement may be +explained by reference to certain modifications of it. It must, however, +be considered as an abstraction, and should be looked on in the light +rather of a scaffolding, which enables us to see the building and its +several parts, than of the edifice itself, but which latter, from our +imperfect knowledge and limited powers, we could not see without some +such assistance. + +The typical form may be, hypothetically at least, considered as the +primitive one transmitted by hereditary descent from generation to +generation, and modified to suit the requirements of the individual, or +in accordance with circumstances. If it be borne in mind that it is but +an artificial contrivance, more or less true--a means to an end, and not +the end itself--no harm will arise from its employment; and as knowledge +increases, or as circumstances demand, the hypothetical type can be +replaced by another more in accordance with the actual state of science. + + +Teratological changes in the arrangement of organs depend upon arrest of +growth, as when parts usually spirally arranged remain verticillate, +owing to the non-development of the internodes, or to excessive growth, +or development; but in many instances it is impossible, without +studying the development of the malformed flower, to ascertain whether +the altered arrangement is due to an excessive or to a diminished +action. Practically, however, it is of comparatively little importance +to know whether, say, the isolation of parts, that are usually combined +together, is congenital (_i.e._ the result of an arrest of growth +preventing their union), or whether it be due to a separation of parts +primitively undivided; the effect remains the same, though the cause may +have been very different. + +The principal alterations to be mentioned under this head may therefore +be conveniently arranged under the following categories:--Union, +Independence, Displacement, Prolification, Heterotaxy, and Heterogamy. + + + + +PART I. + +UNION OF ORGANS. + + +The union of parts, usually separate in their adult condition, is of +very common occurrence as a malformation. The instances of its +manifestation admit of being grouped under the heads of Cohesion, where +parts of the same whorl, or of the same organ, are united together; and +of Adhesion, where the union takes place between members of different +whorls, or between two or more ordinarily wholly detached and distinct +parts. In either case, the apparent union may be congenital (that is, +the result of a primitive integrity or a lack of separation), or it may +really consist in a coalition of parts originally distinct and separate. +In practice it is not always easy to distinguish between these two +different conditions. Indeed, in most cases it cannot be done without +tracing the development of the flower throughout all its stages. It is +needless to make more than a passing allusion to the frequency with +which both congenital integrity or subsequent coalescence of organs +exist under ordinary circumstances. Considered as a teratological +phenomenon, union admits of being grouped into several subdivisions, +such as Cohesion, Adhesion, Synanthy, Syncarpy, Synophty, &c. Each of +these subdivisions will be separately treated, but it maybe here said +that, in all or any case, the degree of fusion may be very slight, or it +may be so perfect that there may be a complete amalgamation of two or +more parts, while to all outward appearance the organ may be single. +The column of Orchids may be referred to as an illustration under +natural circumstances of the complete union of many usually distinct +parts. + +In the uncertainty that exists in many cases as to the real nature of +the occurrence, it would be idle to attempt to explain the causes of +fusions. It is clear, however, that an arrest of development will tend +towards the maintenance of primordial integrity (congenital fusion), and +that pressure will induce the coalition of organs primarily distinct. + + + + +CHAPTER I. + +COHESION. + + +Following Augustin Pyranius De Candolle, botanists have applied the term +cohesion to the coalescence of parts of the same organ or of members of +the same whorl; for instance, to the union of the sepals in a +gamosepalous calyx, or of the petals in a gamopetalous corolla. It may +arise either from a union between organs originally distinct, or more +frequently from a want of separation between parts, which under general +circumstances become divided during their development. Nothing is more +common as a normal occurrence, while viewed as a teratological +phenomenon it is also very frequent. For the purposes of convenience it +admits of subdivision into those cases wherein the union takes place +between the branches of the same plant, or between the margins of the +same leaf-organ, or between those of different members of the same +whorl. + +=Cohesion between the axes of the same plant.=--This cohesion may occur +in various manners. Firstly. The branches of the main stem may become +united one to the other. Secondly. Two or more stems become joined +together. Thirdly. The branches become united to the stem; or, lastly, +the roots may become fused one with another. + +[Illustration: FIG. 2--Cohesion of two branches in _Dipsacus +sylvestris_.] + +The first of these is most commonly met with, doubtless owing to the +number of the branches and the facilities for their union. An +illustration of it is afforded by the figure (fig. 2), showing cohesion +affecting the branches of a teazle (_Dipsacus sylvestris_). Union of the +branches may be the result of an original cohesion of the buds, while in +other cases the fusion does not take place until after development has +proceeded to some extent. Of this latter kind illustrations are common +where the branches are in close approximation; if the bark be removed by +friction the two surfaces are very likely to become united (natural +grafting). Such a union of the branches is very common in the ivy, the +elder, the beech, and other plants. It may take place in various +directions, lengthwise, obliquely, or transversely, according to +circumstances. This mode of union belongs, perhaps, rather to the domain +of pathology than of teratology. Some of the instances that have been +recorded of very large trees, such as the chestnut of Mount AEtna, are +really cases where fusion has taken place between several of the +branches, or suckers, thrown out from the same original stem.[10] The +same process of grafting occurs sometimes in the roots, as in _Taxus +baccata_ mentioned by Moquin, and also in the aerial roots of many of +the tropical climbing plants, such as _Clusia rosea_, &c. + +[Illustration: FIG. 3.--Fasciation in Lettuce.] + +=Fasciation.=--In the preceding instances of union between the branches, +&c., the actual number of the fused parts is not increased; but if it +happen that an unusual number of buds be formed in close apposition, so +that they are liable to be compressed during their growth, union is very +likely to take place, the more so from the softness of the young +tissues. In this way it is probable that what is termed fasciation is +brought about. This is one of the most common of all malformations, and +seems to affect certain plants more frequently than others. In its +simplest form it consists of a flat, ribbon-like expansion of the stem +or branch; cylindrical below, the branches gradually lose their pristine +form, and assume the flattened condition. + +[Illustration: FIG. 4.--Fasciation in _Asparagus_.] + +[Illustration: FIG. 5.--Fasciated branch of _Pinus Pinaster_.] + +Very generally the surface is striated by the prominence of the woody +fibres which, running parallel for a time, converge or diverge at the +summit according to the shape of the branch. If the rate of growth be +equal, or nearly so, on both sides, the stem retains its straight +direction, but it more generally happens that the growth on one side is +more rapid and more vigorous than on the other, and hence arises that +curvature of the fasciated branch so commonly met with, _e.g._ in the +ash (_Fraxinus_), wherein it has been likened to a shepherd's crook. It +is probable that almost any plant may present this change. It occurs +alike in herbaceous and in woody plants, originating in the latter case +while the branches are still soft. It may be remarked that, in the case +of herbaceous plants, the fasciation always affects the principal stem, +while, on the other hand, in the case of trees and shrubs the deformity +occurs most frequently in the branches; thus, while in the former it +may be said that the whole of the stem is more or less affected, in the +latter it is rare to see more than one or two branches of the same tree +thus deformed. It is a common thing for the fasciated branch to divide +at the summit into a number of subdivisions. These latter may be +deformed like the parent branch, or they may resume the ordinary aspect +of the twigs. + +[Illustration: FIG. 6.--Fasciation and spiral torsion in the stem of +_Asparagus_.] + +Sometimes the flattened stem is destitute of buds, at other times, these +organs are scattered irregularly over its surface or are crowded +together in a sort of crest along the apex. When, as often happens, the +deformity is accompanied with a twisting of the branch spirally, the +buds may be placed irregularly, or in other cases along the free edge of +the spiral curve. In a specimen of _Bupleurum falcatum_ mentioned by +Moquin the spiral arrangement of the leaves was replaced by a series of +perfect whorls, each consisting of five, six, seven, or eight segments, +and there was a flower-stalk in the axil of each leaf. + +When flowers are borne on these fasciated stems they are generally +altered in structure; sometimes the thalamus itself becomes more or less +fasciated or flattened, and the different organs of the flower are +arranged on an elliptical axis. A case of this nature is described by +Schlechtendal ('Bot. Zeit.,' 1857, p. 880), in _Cytisus nigricans_, and +M. Moquin-Tandon describes an instance in the vine in one flower of +which sepals, petals, stamens, and ovary were abortive, while the +receptacle was hypertrophied and fasciated, and bore on its surface a +few adventitious buds.[11] The pedicels of _Streptocarpus Rexii_ have +also been observed in a fasciated state.[12] + +It has been occasionally observed that the fasciated condition is +hereditary; thus, Moquin relates that some seeds of a fasciated +_Cirsium_ reproduced the same condition in the seedlings,[13] while a +similar tendency is inherited in the case of the cockscomb (_Celosia_). + +With reference to the nature of the deformity in question there is a +difference of opinion; while most authors consider it to be due to the +causes before mentioned, Moquin was of opinion that fasciation was due +to a flattening of a single stem or branch. Linnaeus, on the other hand, +considered such stems to be the result of the formation of an unusual +number of buds, the shoots resulting from which became coherent as +growth proceeded:--"_Fasciata dici solet planta cum plures caules +connascuntur, ut unus ex plurimis instar fasciae evadat et compressus_" +(Linn., 'Phil. Bot.,' 274). A similar opinion was held by J. D. Major in +a singular book entitled 'De Planta, Monstrosa, Gottorpiensi,' +Schleswig, 1665, wherein the stem of a _Chrysanthemum_ is depicted in +the fasciated condition. + +[Illustration: FIG. 7.--Fasciation in the scape of the Dandelion +(_Leontodon Taraxacum_).] + +The striae, which these stems almost invariably present, exhibit the +lines of junction, and the spiral or other curvatures and contraction, +which are so often met with, may be accounted for by the unequal growth +of one portion of the stem as contrasted with that of another. Against +this view Moquin cites the instances of one-stemmed plants, such as +_Androsace maxima_, but, on the other hand, those herbaceous plants +having usually but a single stem not unfrequently produce several which +may remain distinct, but not uncommonly become united together. Prof. +Hincks[14] cites cases of this kind in _Primula vulgaris_, _Hieracium +aureum_, and _Ranunculus bulbosus_. I have myself met with several cases +of the kind in _Primula veris_, in the Polyanthus, in the Daisy, and in +the _Leontodon Taraxacum_, in which latter a fusion of two or more +flower-stems bearing at the top a composite flower, and made up of two, +three, four, or more flowers combined together, and containing all the +organs that would be present in the same flowers if separate, is very +common. + +Moquin's second objection is founded upon the fact that, in certain +fasciated stems, the branches are not increased in number or altered in +arrangement from what is usual; but however true this may be in +particular cases, it is quite certain that in the majority of instances +a large increase in the number of leaves and buds is a prominent +characteristic of fasciated stems. + +Another argument used by the distinguished French botanist to show that +fasciated stems are not due to cohesion of two or more stems, is founded +on the fact that a transverse section of a fasciated stem generally +shows an elliptical outline with but a single central canal. On the +other hand, if two branches become united and a transverse section be +made, the form of the cut surface would be more or less like that of the +figure 8[Symbol: 8 turned 90 deg.], although in old stems this may give +place to an elliptical outline, but even then traces of two medullary +canals may be found. This argument is very deceptive, for the appearance +of the transverse section must depend, not only on the intimacy of their +union, but also on the internal structure of the stems themselves. When +two flowers cohere without much pressure they exhibit uniting circles +somewhat resembling the figure of 8[Symbol: 8 turned 90 deg.], but when +more completely combined they have an outline of a very elongated figure, +and something similar is to be expected in herbaceous stems. Even the +elongated pith of a transversely cut, woody, fasciated stem only marks +the intimate union of several branches, and Prof. Hincks, whose views +the writer entirely shares, has noticed instances of the union of two, +and of only two, stems where the internal appearance was the same as in +other fasciations. + +Moquin, moreover, raises the objection that it is unlikely that several +branches should become united lengthwise in one plane only, and, +further, that in the greater number of fasciations all the other +branches which should be present are to be found--not one is wanting, +not one has disappeared, as might have been anticipated had fusion taken +place. In raising this objection, Moquin seems not sufficiently to have +considered the circumstance that the buds in these cases are in one +plane from the first, and are all about equal in point of age and size. + +The last objection that Moquin raises to the opinion that fasciation is +the result of a grafting process is, that in such a case, examples +should be found wherein the branches are incompletely fused, and where +on a transverse section traces of the medullary canals belonging to each +branch should be visible. The arrangement of leaves or buds on the +surface should also in such a case indicate a fusion of several spiral +cycles or whorls. To this it may be replied that such cases are met with +very frequently indeed. A figure is given by De Candolle[15] of a stem +of _Spartium junceum_ having several branches only imperfectly +fasciated. + +Fasciated stems, then, seem to be best explained, as is stated by Prof. +Hincks, "on the principle of adhesion arising in cases where from +superabundant nourishment, especially if accompanied by some check or +injury, numerous buds have been produced in close proximity, and the +supposition that these growths are produced by the dilatation of a +single stem is founded on a false analogy between fasciated stems and +certain other anomalous growths." + +It will not, of course, be forgotten that this fasciated condition +occurs so frequently in some plants as almost to constitute their +natural state, _e.g._ _Sedum cristatum_, _Celosia_, &c. This condition +may be induced by the art of the gardener--"_Fit idem arte, si plures +caules enascentes cogantur penetrare coarctatum spatium et parturiri +tanquam ex angusto utero, sic saepe in Ranunculo, Beta, Asparago, +Hesperide Pinu, Celosia, Tragopogone, Scorzonera Cotula foetida_," +Linnaeus op. cit. + +Plot, in his 'History of Oxfordshire,' considers fasciation to arise +from the ascent of too much nourishment for one stalk and not enough for +two, "which accident of plants," says Plot, the German virtuosi ('Misc. +Curios. Med. Physic. Acad. Nat. Cur.,' Ann. i, Observ. 102,) "think only +to happen after hard and late winters, by reason whereof, indeed, the +sap, being restrained somewhat longer than ordinary, upon sudden thaws +may probably be sent up more forcibly, and so produce these fasciated +stalks, whereas the natural and graduated ascent would have produced +them but single." Prof. Hincks' explanation is, however, more near to +the truth, and his opinion is borne out by the frequency with which this +change is met with in certain plants which are frequently forced on +during their growth, as lettuce, asparagus, endive, &c., all of which +are very subject to this change. In the 'Transactions of the +Horticultural Society of London,' vol. iv, p. 321, Mr. Knight gives an +account of the cultivation of the cockscomb, so as to ensure the +production of the very large flower-stalks for which this plant is +admired. The principal points in the culture were the application of a +large quantity of stimulating manure and the maintenance of a high +temperature. One of them so grown measured eighteen inches in width. + +The list which is appended is intended to show those plants in which +fasciation has been most frequently observed. It makes no pretension to +be complete, but is sufficiently so for the purpose indicated: the * +denotes the especial frequency of the change in question; the ! +indicates that the writer has himself seen the plant, so marked, +affected in this way. The remainder have been copied from various +sources. + + EXOGENS. + + [Greek: alpha]. _Herbaceous._ + + Ranunculus tripartitus. + * bulbosus! + Philonotis. + Delphinium elatum. + * sp.! + Hesperis matronalis. + *Cheiranthus Cheiri! + *Matthiola incana! + *Brassica oleracea! var. pl. inflor. + Linum usitatissimum! + Althaea rosea! + Lavatera trimestris. + Geranii sp. + Tropaeolum majus! + Viola odorata inflor.! + Reseda odorata! + Fragaria vesca. + Ervum lens. + Trifolium resupinatum. + repens! + pratense! + Saxifraga mutata. + irrigua. + Bupleurum falcatum. + Bunium flexuosum. + *Sedum reflexum! + cristatum! + Epilobium augustifolium! + Momordica Elaterium! + Gaura biennis. + Cotula foetida. + Barkhausia taraxacifolia. + Carlina vulgaris! + Apargia autumnalis. + *Leontodon Taraxacum inflor.! + Centaurea Scabiosa. + *Cichorium Intybus! + Hieracium Pilosella. + aureum. + umbellatum. + *Chrysanthemum Leucanthemum. + indicum! + Anthemis nobilis. + arvensis. + Cirsium lanceolatum. + Conyza squarrosa! + Inula dysenterica! + Tragopogon porrifolium. + Cnicus palustris. + Carduus arvensis! + Helianthus tuberosus! + annuus. + Cineraria palustris. + Helianthus sp.! + Dahlia variabilis. + Bellis perennis inflor.! + Coreopsis sp.! + Crepis virens. + Lactuca sativa! + Zinnia elegans. + *Campanula medium! + rapunculoides. + thyrsoidea. + Dipsacus pilosus. + fullonum. + silvestris. + Knautia arvensis. + Phyteuma orbiculare. + Jasione montana. + *Linaria purpurea! + Antirrhinum majus! + Veronica amethystea. + Veronica maritima. + sp. + Russellia juncea! + Digitalis purpurea! + Ajuga pyramidalis. + Hyssopus officinalis. + Dracocephalum moldavicum. + Myosotis scorpioides. + Echium pyrenaicum. + simplex. + Stapeliae sp. + Lysimachia vulgaris! + Androsace maxima. + Primula veris inflor.! + denticulata inflor.! + Polemonium coeruleum. + Convolvulus sepium! + arvensis! + Plantago media. + *Euphorbia Characias. + exigua. + * Cyparissias. + Suaeda maritima. + *Celosia sp. + Beta vulgaris inflor.! + Phytolacca sp. + + [Greek: beta]. _Woody._ + + Berberis vulgaris. + Hibiscus syriacus! + Acer pseudo-platanus! + Dodonaea viscosa. + Sterculia platanifolia. + Euonymus japonicus! + Vitis vinifera inflor.! + Spartium Scoparium! + Spartium junceum! + Cytisus Laburnum. + nigricans. + Chorozema ilicifolium. + Amorpha sp. + Phaseolus sp. + Prunus sylvestris. + Laurocerasus! + Rosa sp.! + Spiraea sp.! + Cotoneaster microphylla! + Ailanthus glandulosus. + *Fraxinus Ornus! + * excelsior! + Melia Azedarach. + Xanthoxylum sp.! + Sambucus nigra.! + Aucuba japonica. + Erica sp. cult. + Jasminum nudiflorum! + officinale! + Olea europoea. + Punica Granatum. + Ilex aquifolium! + Daphne indica. + Daphne odora. + Suaeda fruticosa. + Ulmus campestris. + Alnus incana. + Salix vitellina, &c.! + Thuja orientalis. + Pinus pinaster! + sylvestris! + Abies excelsa! + Taxus baccata. + Larix europoea. + + ENDOGENS. + + Lilium Martagon. + candidum! + *Fritillaria imperialis! + Asparagus officinalis! + Hyacinthus orientalis! + Tamus communis! + Narcissi sp.! + Gladiolus sp. + Zea Mays. + Filices. + + See also--Moquin-Tandon, 'Elem. Ter. Veget.,' p. 146; C. O. + Weber, 'Verhandl. Nat. Hist.,' Vereins, f. d. Preuss., Rheinl. + und Westphal., 1860, p. 347, tab. vii; Hallier, 'Phytopathol.,' + p. 128; Boehmer, 'De plantis Fasciatis,' Wittenb., 1752. + +=Cohesion of foliar organs.=--This takes place in several ways, and in +very various degrees; the simplest case is that characterised by the +cohesion of the margins of the same organ, as in the condition called +perfoliate in descriptive works, and which is due either to a cohesion +of the margins of the basal lobes of the leaf, or to the development of +the leaf in a sheathing or tubular manner. As an abnormal occurrence, I +have met with this perfoliation in a leaf of _Goodenia ovata_. The +condition in question is often loosely confounded with connation, or the +union of two leaves by their bases. In other cases the union takes place +between the margins of two or more leaves. + +=Cohesion of margins of single organs.=--The leaves of Hazels may often +be found with their margins coherent at the base, so as to become +peltate, while in other cases, the disc of the leaf is so depressed that +a true pitcher is formed. This happens also in the Lime _Tilia_, in +which genus pitcher- or hood-like leaves (_folia cucullata_) may +frequently be met with. There are trees with leaves of this character in +the cemetery of a Cistercian Monastery at Sedlitz, on which it is said +that certain monks were once hung: hence the legend has arisen, that the +peculiar form of the leaf was given in order to perpetuate the memory of +the martyred monks. ('Bayer. Monogr. _Tiliae_,' Berlin, 1861.) It is also +stated that this condition is not perpetuated by grafting. + +[Illustration: FIG. 8.--Pitcher-shaped leaf of _Pelargonium_.] + +I have in my possession a leaf of _Antirrhinum majus_, and also a +specimen of _Pelargonium_, wherein the blade of the leaf is funnel-like, +and the petiole is cylindrical, not compressed, and grooved on the upper +surface, as is usually the case. A comparison of the leaves of +_Pelargonium peltatum_ with those of _P. cucullatum_ ('Cav. Diss.,' +tab., 106) will show how easy the passage is from a peltate to a tubular +leaf. In these cases the tubular form may rather be due to dilatation +than to cohesion. M. Kickx[16] mentions an instance of the kind in the +leaves of a species of _Nicotiana_, and also figures the leaf of a rose +in which two opposite leaflets presented themselves in the form of +stalked cups. Schlechtendal[17] notices something of the same kind in +the leaf of _Amorpha fruticosa_; Treviranus[18] in that of _Aristolochia +Sipho_. + +M. Puel[19] describes a leaf of _Polygonatum multiflorum_, the margins +of which were so completely united together, as only to leave a circular +aperture at the top, through which passed the ends of the leaves. The +Rev. Mr. Hincks, at the meeting of the British Association at Newcastle +(1838), showed a leaf of a Tulip, whose margins were so united that the +whole leaf served as a hood, and was carried upwards by the growing +flower like the calyptra of a Moss. + +The margins of the stipules are also occasionally united, so as to form +a little horn-shaped tube. I have met with instances of this kind in the +common white clover, _Trifolium repens_, where on each side of the base +of the petiole the stipules had the form just indicated. That the bracts +also may assume this condition, may be inferred from the peculiar +horn-like structures of _Marcgraavia_, which appear to originate from the +union of the margins of the reflected leaf. + +=Tubular petals= occur normally in some flowers, as _Helleborus_, +_Epimedium_, _Viola_, &c., and as an exceptional occurrence I have seen +them in _Ranunculus repens_, while in _Eranthis hyemalis_ transitions +may frequently be seen between the flat outer segments of the perianth +and the tubular petals. To Dr. Sankey, of Sandywell Park, I am indebted +for the flower of a Pelargonium, in which one of the petals had the +form of a cup supported on a long stalk. This cup-shaped organ was +placed at the back of the flower, and had the dark colour proper to the +petals in that situation. I have seen a petal of Clarkia similarly +tubular, while some of the cultivated varieties of _Primula sinensis_ +exhibit tubular petals so perfect in shape as closely to resemble +perfect corollas. + +[Illustration: FIG. 9.--_Eranthis hyemalis_. Transition from flat sepal +to tubular petal.] + +Like the petals, the stamens, and even the styles, assume a hollow +tubular form. This change of form in the case of the stamens is, of +course, usually attended by the petaloid expansion of the filament, or +anther, and the more or less complete obliteration of the pollen sacs, +as in Fuchsias, and in some double-flowered Antirrhinums.[20] So also in +some semi-double varieties of _Narcissus poeticus_, and in _Aquilegia_. +By the late Professor Charles Morren, this affection of the stamens and +pistils was called _Solenaidie_,[21] but as a similar condition exists +in other organs, it hardly seems worth while to adopt a special term for +the phenomenon, as it presents itself in one set of organs. + +In many of these cases it is difficult to say whether the cup-like or +tubular form is due to a dilatation or hollowing out of the organ +affected, or to a fusion of its edges. The arrangement of the veins will +in some cases supply the clue, and in others the regularity of form +will indicate the nature of the malformation, for in those instances +where the cup is the result of expansion, its margin is more likely to +be regular and even than in those where the hollow form is the result of +fusion. + +=Cohesion of several organs by their margins:--leaves, &c.=--The union +of the margins of two or more different organs is of more common +occurrence than the preceding, the leaves being frequently subjected to +this change. Occasionally, the leaflets of a compound leaf have been +observed united by their margins, as in the strawberry, the white +trefoil, and others. Sometimes the union takes place by means of the +stalks only. I have an instance of this in a Pelargonium, in _Tropaeolum +majus_, and _Strelitzia regina_; in other cases, the whole extent of the +leaf becomes joined to its neighbour, the leaves thus becoming +completely united by their edges, as in those of _Justicia_, +_oxyphylla_.[22] M. Clos[23] has observed the same thing in the leaves +of the lentil _Ervum lens_, conjoined with fasciation of the stem, and +many other examples might be given. Some of the recorded cases are +probably really due to fission of one leaf into two rather than to +fusion. Although usually the lower portions of the leaf are united +together, leaving the upper parts more or less detached, there are some +instances in which the margins of the leaf at their upper portion have +been noticed to be coherent, while their lower portions, with their +stalks, were completely free.[24] + +Cohesion of the leaves frequently accompanies the union of the branches +and fasciation as might have been anticipated. Moquin cites the +fenestrated leaves of _Dracontium pertusum_, as well as some cases of a +similar kind that are occasionally met with, as instances of the +cohesion of the margins at the base and apex of the leaf, which thus +appears perforated. This appearance, however, is probably due to some +other cause. When the leaves are verticillate and numerous, and they +become coherent by their margins, they form a foliaceous tube around the +stem. When there are but two opposite leaves, and these become united by +their margins, we have a state of things precisely resembling that to +which the term connate is applied. + +Fusion of the edges of the cotyledons also occasionally takes place, as +in _Ebenus cretica_.[25] It has also been observed in _Tithonia_, and is +of constant occurrence in the seed leaves of some _Mesembryanthema_. +This condition must be carefully distinguished from the very similar +appearance produced by quite a different cause, viz., the splitting of +one cotyledon into two, which gives rise to the appearance as if two +were partially united together. + +Some of the ascidia or pitcher-like formations are due to the cohesion +of the margins of two leaves, as in a specimen of _Crassula +arborescens_, observed by C. Morren. + +[Illustration: FIG. 10.--Two-leaved pitcher of _Crassula arborescens_, +after C. Morren.] + +The stipules may also be fused together in different ways; their edges +sometimes cohere between the leaf and the stem, and thus form a solitary +intra-axillary stipule. At other times they become united in such a +manner as to produce a single notched stipule opposite to the leaf. +Again, in other cases, they are so united on each side of the stem, that +in place of four there seem only to exist two, common to the two leaves +as in the Hop. + +To the Rev. M. J. Berkeley I am indebted for specimens of a curious +pitcher-like formation in the garden Pea. The structure in question +consisted of a stalked foliaceous cup proceeding from the inflorescence. +On examination of the ordinary inflorescence, there will be seen at the +base of the upper of two flowers a small rudimentary bract, having a +swollen circular or ring-like base, from which proceeds a small +awl-shaped process, representing the midrib of an abortive leaf. In some +of Mr. Berkeley's specimens, the stipules were developed as leafy +appendages at the base of the leaf-stalk or midrib, the latter retaining +its shortened form, while, in others, the two stipules had become +connate into a cup, and all trace of the midrib was lost. The cup in +question would thus seem to have been formed from the connation of two +stipules which are ordinarily abortive. + +Cohesion of the bracts by their edges, so as to form a tubular +involucre, or by their surfaces, so as to form a cupule, is not of +uncommon occurrence, under natural conditions, and may be met with in +plants which ordinarily do not exhibit this appearance. + +=Cohesion of the sepals= in a normally polypetalous calyx renders the +latter gamosepalous, and is not of uncommon occurrence, to a partial +extent, though rarely met with complete. I have observed a junction of +the sepals to be one of the commonest malformations among Orchids, +indeed such a state of things occurs normally in _Masdevallia +Cypripedium_, &c. An illustration of this occurrence is given by Mr. J. +T. Moggridge in _Ophrys insectifera_, in 'Seemann's Journal of Botany,' +1866, p. 168, tab. 47. In Orchids, this cohesion of sepals is very often +co-existent with other more important changes, such as absence of the +labellum, dislocation of the parts of the flower, &c. + +[Illustration: FIG. 11.--Gamopetalous flower of _Papaver bracteatum_.] + +=Cohesion of the petals.=--Linnaeus mentions the occurrence of cohesion +of the petals in _Saponaria.[26]_ Moquin notices a Rose in which the +petals were united into a long tube, their upper portions were free and +bent downwards, forming a sort of irregular limb. An instance of the +polypetalous regular perianth of _Clematis viticella_ being changed into +a monopetalous irregular one, like the corolla of Labiates, is recorded +by Jaeger.[27] There is in cultivation a variety of _Papaver +bracteatum_, in which the petals are united by their margins so as to +form a large cup. Under normal circumstances, the petals become fused +together by their edges along their whole extent, at the base only, at +the apex only, as in the Vine, or at the base and apex, leaving the +central portions detached. Indications of the junction of the petals may +generally be traced by the arrangement of the veins, or by the notches +or lobes left by imperfect union. In Crocuses I have frequently met with +cohesion of the segments of the perianth, by means of their surfaces, +but the union was confined to the centre of the segment, leaving the +rest of the surfaces free. + +=Cohesion of the stamens.=--Under natural circumstances, cohesion of the +stamens is said to take place either by the union of their filaments, so +as to form one, two, or more parcels (Monadelphia, Diadelphia, +Polyadelphia); at other times, by the cohesion of the anthers +(Syngenesia), in which latter case the union is generally very slight. +It must be remembered, however, that the so-called cohesion of the +filaments is in many cases due rather to the formation of compound +stamens, _i.e._ to the formation from one original staminal tubercle of +numerous secondary ones, so that the process is rather one of over +development than of fusion or of disjunction. These conditions may be +met with as accidental occurrences in plants or in flowers, not usually +showing this arrangement. Thus, for instance, Professor Andersson, of +Stockholm, describes a monstrosity of _Salix calyculata_, in which the +stamens were so united together as to form a tube open at the top like a +follicle.[28] This is an exaggerated degree of that fusion which exists +normally in _Salix monandra_, in Cucurbits and other plants. + +=Cohesion of the pistils= is also of very frequent occurrence in plants, +under ordinary circumstances, but is less commonly met with than might +have been expected as a teratological phenomenon. + + Further details relating to cohesion of the various parts of + the flower are cited in Moquin-Tandon, 'El. Ter. Veg.,' p. 248; + 'Weber. Verhandl. Nat. Hist. Vereins f. d. Preuss. Rheinl. und + Westphal.,' 1860, p. 332, tabs. 6 et 7. + +=Formation of ascidia or pitchers.=--In the preceding paragraphs, the +formation of tubular or horn-like structures, from the union of the +margins of one organ, or from the coalescence, or it may be from the +want of separation of various organs, has been alluded to, so that it +seems only necessary now, by way of summary, to mention the +classification of ascidia proposed by Professor Charles Morren[29], who +divides the structures in question into two heads, according as they are +formed from one or more leaves. The following list is arranged according +to the views of the Belgian savant, and comprises a few additional +illustrations. Those to which the ! is affixed have been seen by the +writer himself; the * indicates the more frequent occurrence of the +phenomenon in some than in other plants. Those plants, such as +_Nepenthes_, &c., which occur normally and constantly, are not here +included. Possibly some of the cases would be more properly classed +under dilatation or excavation. + + ASCIDIA. + + A. _Monophyllous_. + + 1. Sarracenia-like pitchers, formed by a single leaf, the edges + of which are united for the greater portion of their length, + but are disunited near the top, so as to leave an oblique + aperture. + + *Brassica oleracea (several of the cultivated varieties)! + *Tilia europaea! + Pelargonium inquinans! + Staphylea pinnata. + Amorpha fruticosa. + Pisum sativum! + Lathyrus tuberosus. + Vicia sp. + Gleditschia sp. + Ceratonia siliqua. + Trifolium repens! + Cassia marylandica. + Mimosa Lophantha. + Rosa centifolia. + gallica. + Begonia sp. + Bellis perennis! + Nicotiana sp. + Goodenia ovata! + Antirrhinum majus! + Vinca rosea. + Polygonum orientale. + Aristolochia sipho? + Codiaeum variegatum var.! + Spinacia oleracea. + Corylus avellana! + Polygonatum multiflorum. + Xanthosoma appendiculatum! + + 2. Calyptriform or hood-like pitchers, formed by the complete + union of the margins, and falling off by a transverse fissure + (as in the calyx of Escholtzia). + + Tulipa Gesneriana. + + B. _Polyphyllous._ + + 1. Diphyllous, formed by the union of two leaves into a single + cup, tube, or funnel, &c. + + Pisum sativum (stipules)! + Crassula arborescens. + Polygonatum multiflorum. + + 2. Triphyllous, formed by the union of three leaves. + + Paris quadrifolia var. + + Besides the above varieties of ascidia formed from the union of + one or more leaves, there are others which seem to be the + result of a peculiar excrescence or hypertrophy of the leaf. + Such are some of the curious pitcher-like structures met with + occasionally in the leaves of cabbages, lettuces, Aristolochia, + &c. See Hypertrophy, cup-like deformities, &c. + + In addition to other publications previously mentioned, + reference may be made to the following treatises on the subject + of ascidia:--Bonnet, 'Rech. Us. Feuilles,' p. 216, tab. xxvi, + f. 1, _Brassica_; De Candolle, 'Trans. Hort. Soc.,' t. v, pl. + 1, _Brassica_; Id., 'Org. Veget.,' I, 316; 'Bull. Soc. Bot. + Fr.,' I, p. 62, _Polygonatum_; 'Bull. Acad. Belg.,' 1851, p. + 591, _Rosa_; Hoffmann, 'Tijdschrift v. Natuur. Geschied.,' vol. + viii, p. 318, tab. 9, _Ceratonia_; C. Mulder, 'Tijdschrift, + &c.,' vol. vi, p. 106, tab. 5, 6, _Trifolium_, _Mimosa_, + _Staphylea_;' Molkenboer,' p. 115, t. 4, _Brassica_. + +FOOTNOTES: + +[10] See a curious instance of this kind in the branches of _Pinus_. +'Regel. Garten Flora,' vol. 8, tab. 268. + +[11] 'Bull. Soc. Bot. France,' 1860, p. 881. + +[12] Ibid., 1861, p. 708. + +[13] Ibid., 1860, p. 923. + +[14] 'Proc. Linn. Soc.,' April 5, 1853. + +[15] 'Organ. Veget.,' pl. iii, fig. 1. + +[16] 'Bull. Acad. Roy. Bruxelles,' t. xviii, p. i and p. 591. + +[17] 'Linnaea,' tom. 13, p. 383. + +[18] 'Verhandl. Nat. Hist. Vereins,' 1859, Bonn, tom. xvi, tab. 3. + +[19] 'Bull. Soc. Bot. Fr.,' vol. i, p. 62. + +[20] 'Report of Internat. Bot. Congress,' London, 1866, p. 131, tab. +vii, figs. 10-13. + +[21] 'Bull. Acad. Roy. Belg.,' t. xviii, 2nd part, p. 179. + +[22] D. C., 'Organ. Veget.,' pl. xvii, fig. 3, and pl. xlviii, fig. 2. + +[23] 'Mem. Acad. Toulouse,' 1862. + +[24] Bonnet, 'Recherches Us. feuill.,' pl. xxi, fig. 2. + +[25] De Candolle, 'Mem. Leg.,' pl. v, fig. 14. + +[26] 'Phil. Bot.,' Sec. 125. + +[27] 'Nov. Act. Acad. Nat. Cur.,' 14, p. 642, t. xxxvii. + +[28] 'Journal of the Linn. Soc. Bot.,' vol. iv, p. 55. + +[29] 'Bull. Acad. Roy. Bruxelles,' 1838, t. v, p. 582. 'Bull. Acad. Roy. +Belg.,' 1852, t. xix, part iii, p. 437. + + + + +CHAPTER II. + +ADHESION. + + +Adhesion, so called, occurs either from actual union of originally +distinct members of different whorls or from the non-occurrence of that +separation which usually takes place between them. It is thus in some +degree a graver deviation than cohesion, and is generally a consequence +of, or at least is coexistent with, more serious changes; thus if two +leaves of the same whorl are coherent the change is not very great, but +if two leaves belonging to different whorls, or two leaves in the same +spiral cycle are adherent, a deformation in the axis or a certain amount +of dislocation must almost necessarily exist. Adhesion as a normal +occurrence is usually the result of a lack of separation rather than of +union of parts primitively separate. Instances of adhesion between +different organs is seen under ordinary circumstances in the bract of +the Lime tree, which adheres to the peduncle, also in _Neuropeltis_, +while in _Erythrochiton hypophyllanthus_ the cymose peduncles are +adherent to the under surface of the leaf. + +Adhesion between the axes of the same plant is sufficiently treated of +under the head of Cohesion, from which it is in this instance impossible +to make a distinction. Adhesion of the inflorescence is necessarily a +frequent accompaniment of fasciation and cohesion of the branches. + +=Adhesion of foliar organs= may occur either between the margins or +between the surfaces of the affected parts; in the former case there is +almost necessarily more or less displacement and change of direction, +such as a twisting of the stem and a vertical rather than a horizontal +attachment of the foliar organ to it; hence it generally forms but a +part of other and more important deviations. + +=Adhesion of leaves by their surfaces.=--The union of leaves by their +surfaces is not of very frequent occurrence, many of the instances cited +being truly referable to other conditions. Bonnet describes the union of +two lettuce leaves, and Turpin that of two leaves of _Agave americana_, +in which latter the upper surface of one leaf was adherent to the lower +surface of the leaf next above it, and I have myself met with similar +instances in the wallflower and in lettuce and cabbage leaves; other +instances have been mentioned in _Saxifraga_, _Gesnera_, _&c._[30] + +In these cases, owing to the non-development of the internodes, the +nascent leaves are closely packed, and the conditions for adhesion are +favorable, but in most of the so-called cases of adhesion of leaf to +leaf by the surface, a preferable explanation is afforded either by an +exuberant development (hypertrophy) or by chorisis (see sections on +those subjects). Thus, when a leaf of this kind is apparently so united, +that the lower surface of one is adherent to the corresponding surface +of another, the phenomenon is probably due rather to extra development +or to fission. There is an exception to this, however, in the case of +two vertically-erect leaves on opposite sides of the stem; here the two +upper or inner surfaces may become adherent, as in an orange, where two +leaves were thus united, the terminal bud between them being suppressed +or abortive. + +Adhesion between the membranous bract of _Narcissus poeticus_ and the +upper surface of the leaf is described by Moquin.[31] The same author +mentions having seen a remarkable example of adhesion in the involucels +of _Caucalis leptophylla_, the bracts of which were soldered to the +outer surface of the flowers. M. Bureau[32] mentions an instance +wherein the spathe of _Narcissus biflorus_ was partially twisted in such +a manner that the lower surface of its median nerve was adherent to the +corresponding surface of one of the sepals, mid-rib to mid-rib, thus +apparently confirming a law of G. de Hilaire, that when two parts of the +same individual unite, they generally do so by the corresponding +surfaces or edges, but the rule is probably not so general in its +application as has been supposed. + +=Adhesion of foliar to axile organs.=--The appendicular organs may +likewise be found united to the axile ones. This union takes place in +many ways; sometimes the leaves do not become detached from the stem for +a considerable distance, as in the so-called decurrent leaves, at other +times the leaves are prolonged at their base into lobes, which are +directed along the stem, and are united with it. Turpin records a +tendril of a vine which was fused with the stem for some distance, and +bore leaves and other tendrils. Union of the leaf or bract with the +flower-stalk is not uncommon. It occurs normally in the Lime and other +plants. + +=Adhesion of the sepals to the petals= is spoken of by Morren as +calyphyomy, [Greek: kalyx phyomai.][33] Moquin cites an instance in +_Geranium nodosum_, in which one petal was united by its lower surface +to one of the segments of the calyx. A similar circumstance has been +observed in _Petunia violacea_ by Morren. Duchartre describes an +instance wherein one of the outer sepals of _Cattleya Forbesii_ was +adherent to the labellum.[34] + +=Adhesion of the stamens to the petals= is of common occurrence under +natural circumstances. Cassini has described a malformation of +_Centaurea collina_, in which two of the five stamens were completely +grafted with the corolla, the three others remaining perfectly free. +Adhesion of the petals to the column is not of infrequent occurrence +among Orchids. I have observed cases of the adhesion of the segments of +the perianth to the stamen in _Ophrys aranifera_, _Odontoglossum_, _sp_. +&c. It is the ordinary condition in _Gongora_ and some other genera. I +have seen it also in _Lilium lancifolium_. Some forms of _Crocus_, +occasionally met with, present a very singular appearance, owing to the +adhesion of the stamens to the outer segments of the perianth, the +former, moreover, being partially petaloid in aspect. M. de la Vaud[35] +speaks of a similar union in _Tigridia pavonia_. Morren[36] describes a +malformation of _Fuchsia_ wherein the petals were so completely adherent +to the stamens, that the former were dragged out of their ordinary +position, so as to become opposite to the sepals; the fusion was here so +complete that, no trace of it could be seen externally. It should be +remarked that it was the outer series of stamens that were thus +fused.[37] + +[Illustration: FIG. 12.--_Crocus._ Adhesion of petaloid stamens to +perianth.] + +=Adhesion of stamens to pistils.=--The stamens also may be united to the +pistils, as in gynandrous plants. Moquin speaks of such a case in a +_Scabious_; M. Clos in _Verbascum australe_.[38] I have seen cases of +the same kind in the Wallflower, Cowslip (_Primula veris_), Tulip, +Orange, in the garden Azalea and other plants. + +=Miscellaneous adhesions.=--Sometimes organs, comparatively speaking, +widely separated one from the other, become united together. Miquel has +recorded the union of a stigma with the middle lobe of the lower lip of +the corolla of _Salvia pratensis_.[39] In the accompanying figure [fig. +13], taken from a double wallflower, there is shown an adhesion between +a petal and an open carpel on the one side, and a stamen on the other. + +Moquin speaks of some pears, which were united, at an early stage, with +one or two small leaves borne by the peduncle and grafted to the fruit +by the whole of their upper surface. As the pear increased in size the +leaves became detached from it, leaving on the surface of the fruit an +impression of the same form as the leaf, and differing in colour from +the rest of the surface of the fruit. Traces of the principal nerves +were seen on the pear. + +[Illustration: FIG. 13.--_Cheiranthus cheiri_. Adhesion of petal to +stamen and open carpel.] + +It is curious to notice how very rare it is for the calyx to adhere to +the ovary in flowers where that organ is normally superior. The "_calyx +inferus_" seems scarcely ever to become "_calyx superus_," while, on the +other hand, the "_calyx normaliter superus_" frequently becomes inferior +from detachment from, or from want of union with the surface of the +ovary. + +=Adhesion of fruit to branch.=--Of this Mr. Berkeley[40] cites an +instance in a vegetable marrow (_Cucumis_), where a female flower had +become confluent with the branch, at whose base it was placed, and also +with two or more flowers at the upper part of the same branch, so as to +make an oblique scar running down from the apex of the fruit to the +branch. + +=Synanthy.=--Adhesion of two or more flowers takes place in various +ways; sometimes merely the stalks are united together, so that we have +a single peduncle, bearing at its extremity two flowers placed in +approximation very slightly adherent one to the other. In this manner I +have seen three flowers of the vegetable marrow on a common stalk, the +flowers themselves being only united at the extreme base. Occasionally +cases may be met with wherein the pedicels of a stalked flower become +adherent to the side of a sessile flower. I have noticed this commonly +in _Umbelliferae_. Union of this kind occurs frequently in the common +cornel (_Cornus_), wherein one of the lower flowers becomes adherent to +one of the upper ones. In De Candolle's 'Organographie Vegetale,' Plates +14 and 15, are figured cases of fusion of the flower stems of the +Hyacinth and of a _Centaurea_. In other cases the union involves not +only the stalk but the flowers themselves; thus fusion of the flowers is +a common accompaniment of fasciation, as was the case in the _Campanula_ +figured in the cut (fig. 14). + +[Illustration: FIG. 14.--Synanthic flowers of _Campanula medium_.] + +Synanthy may take place without much derangement of the structure of +either flower, or the union may be attended with abortion or suppression +of some of the parts of one or both flowers. Occasionally this union is +carried to such an extent that a bloom appears to be single, when it is, +in reality, composed of two or more, the parts of which have become not +only fused, but, as it were, thrust into and completely incorporated one +with another, and in such a manner as to occupy the place of some parts +of the flower which have been suppressed. It must not be overlooked that +this adhesion of one flower to another is a very common occurrence under +natural circumstances, as in _Lonicera_, in the common tomato, in +_Pomax_, _Opercularia_, _Symphyomyrtus_, &c., while the large size of +some of the cultivated sunflowers is in like manner due to the union of +two or more flower-heads. + +One of the simplest instances of synanthy is that mentioned by M. +Duchartre,[41] in which two flowers of a hyacinth were united together +simply by means of two segments of the perianth one from each flower. A +similar occurrence has been cited by M. Gay in _Narcissus chrysanthus_. +In like manner the blossoms of Fuchsias or Loniceras occasionally become +adherent merely by their surface, without involving any other change in +the conformation of the flowers. M. Maugin alludes to a case of this +kind in _Aristolochia Clematitis_.[42] + +But it is more usual for some of the organs to be suppressed, so that +the number of existing parts is less than would be the case in two or +more uncombined flowers. A few illustrations will exemplify this. In two +flowers of _Matthiola incana_, that I observed to be joined together, +there were eight sepals, eight petals, and ten perfect stamens, eight +long and two short, instead of twelve. Closer examination showed that +the point of union between the two flowers occurred just where, under +ordinary circumstances, the two short stamens would be. In this instance +but little suppression had occurred. In similar flowers of _Narcissus +incomparabilis_ I remarked a ten-parted perianth, ten stamens within a +single cup, two styles, and a five-celled ovary. Here, then, it would +appear that two segments of the perianth, two stamens, and one carpel +were suppressed. In a Polyanthus there were nine sepals, nine petals, +nine stamens, and a double ovary. + +[Illustration: FIG. 15.--Union of three flowers of _Calanthe vestita_.] + +[Illustration: FIG. 16.--Shows the abortion of the central spur in +synanthic flowers of _Calanthe vestita_.] + +As an illustration of a more complicated nature reference may be made to +three flowers of _Aconitum Napellus_, figured by A. de Chamisso, +'Linnaea,' vol. vii, 1832, p. 205, tab. vii, figs. 1, 2. In this specimen +the two outer blossoms had each four sepals present, namely, the upper +hooded one, one of the lateral sepals, and both of the inferior ones; +the central flower had only the upper sepal and one other, probably one +of the lower sepals; thus there were but ten sepals instead of fifteen. +The nectary-like petals, the stamens, and pistils were all present in +the lateral flowers, but were completely suppressed in the middle one. A +less degree of suppression was exemplified in a triple flower of +_Calanthe vestita_ sent me by Dr. Moore, of Glasnevin, in which all the +parts usually existing in three separate flowers were to be found, with +the exception of the spur belonging to the labellum of the middle flower +(figs. 15, 16). + +One of the most common malformations in the Foxglove (_Digitalis_) +results from the fusion of several of the terminal flowers into one. In +these cases the number of parts is very variable in different instances; +the sepals are more or less blended together, and the corollas as well +as the stamens are usually free and distinct, the latter often of equal +length, so that the blossom, although truly complex, is, as to its +external form, less irregular than under natural circumstances. The +centre of these flowers is occupied by a two to five-celled pistil, +between the carpels of which, not unfrequently, the stem of the plant +projects, bearing on its sides bracts and rudimentary flowers. (See +Prolification.) An instance of this nature is figured in the 'Gardeners' +Chronicle,' 1850, p. 435, from which the cut (fig. 17) is borrowed. + +[Illustration: FIG. 17.--Synanthy and other changes in a Foxglove.] + +One of the most singular recorded instances of changes connected with +fusion of the flowers is that cited by Reinsch,[43] where two female +flowers of _Salix cinerea_ were so united with a male one as to produce +an hermaphrodite blossom. + +It follows, from what has been said, that the number of parts that are +met with in these fused flowers varies according to the number of +blossoms and of the organs which have been suppressed. Comparatively +rarely do we find all the organs present; but when two flowers are +united together we find every possible variety between the number of +parts naturally belonging to the two flowers and that belonging to a +single one. Sometimes instances are met with wherein the calyx does not +present the normal number of parts, while the other parts of the flower +are in excess. I have seen in a _Calceolaria_ a single calyx, with the +ordinary number of sepals, enclosing two corollas, adherent simply by +their upper lips, and containing stamens and pistils in the usual way. +In this instance, then, the sepals of one flower must have been +suppressed, while no such suppression took place in the other parts of +the flower. + +Professor Charles Morren paid special attention to the various methods +in which the flowers of Calceolarias may become fused, and to the +complications that ensue from the suppression of some parts, the +complete amalgamation of others, &c. Referring the reader to the Belgian +savant's papers for the full details of the changes observed, it is only +necessary to allude to a few of the most salient features. + +[Illustration: FIG. 18.--Synanthic flowers of Calceolaria in which, with +two upper lips, there was but a single lower one.] + +Sometimes the upper lips of two flowers are fused into one, the two +lower remaining distinct. In other cases, the upper lip disappears +altogether, while there are two lower lips placed opposite one another;, +of the stamens, sometimes the outermost, at other times the innermost +disappear.[44] + +Occasionally there appears to be, as it were, a transference of the +parts of one flower to another. One of the simplest and most +intelligible cases of this kind is recorded by Wigand in the 'Flora' for +1856, in a compound flower of _Polygonatum anceps_, in which within a +twelve-parted perianth there were twelve stamens and two pistils, one +four-celled, the other two-celled; hence it would appear as if a carpel +belonging to one flower had become united to those constituting the +pistil of the adjacent one. Among Orchids this fusion of some of the +elements of different flowers, together with the suppression of others, +is carried to such an extent as to render the real structure difficult +to decipher. Sometimes flowers of _Ophrys aranifera_, at first sight +seeming normal as to the number, and almost so as regards the +arrangement of their parts, have yet, on examination, proved to be the +result of a confluence of two flowers. Mr. Moggridge has observed +similar phenomena in the same species at Mentone. + +Sometimes the fusion affects flowers belonging to different branches of +the same inflorescence, as in _Centranthus ruber_, described by +Buchenau, 'Flora,' 1857, p. 293, and even a blossom of one generation of +axes may be united with a flower belonging to another generation. Thus +M. Michalet[45] speaks of a case wherein the terminal flower of +_Betonica alopecuros_ was affected with Peloria, and fused with an +adjacent one belonging to a secondary axis of inflorescence, and not yet +expanded. This latter flower had no calyx, but in its place were three +bracts, surrounding the corolla; this again was united to the calyx of +the terminal bloom in a most singular manner, the limb of the corolla +and that of the calyx being so joined one to the other as to form but a +single tube. It is not uncommon, as has been before stated, to find two +corollas enclosed within one calyx, but this is probably the only +recorded instance of the fusion of the calyx and corolla of two +different flowers belonging to two different axes. + +From the preceding details, as well as from others which it is not +necessary to give in this place, it would appear that synanthy is more +liable to occur where the flowers are naturally crowded together[46] +than where they are remote; so too, the upper or younger portions of the +inflorescence are those most subject to this change. In like manner the +derangements consequent on the coalescence of flowers are often more +grave in the central organs, which are most exposed to pressure, and +have the least opportunities of resisting the effects of that agency, +than they are in the outer portions of the flowers where growth is less +restricted. + +Morren in his papers on synanthic _Calceolarias_, before referred to, +considers that the direction in which fusion acts is centripetal, _e.g._ +from the circumference towards the centre of the flower, thus reversing +the natural order of things. He considers that there is a radical +antagonism between the normal organizing forces and the teratological +disorganizing forces, and explains in this way the frequent sterility of +monsters from an imperfect formation of stamens, or pistils, or both. + +The greater tendency in synanthic flowers of parts of one whorl to +adhere to the corresponding organs in another flower has often been +remarked, though the dislocation of parts may be so great as to prevent +this from being carried out in all cases. It appears also that synanthy +is more frequently met with among flowers which have an inferior ovary +than in those in which the relative position of the organ in question +is reversed. This remark applies particularly to individual cases; the +proportion as regards the genera may not be so large. The explanation of +this must of course depend on the circumstances of each particular case; +and it would be wrong to attempt to lay down a general rule, when +organogenists have not yet fully decided in what plants the inferior +ovary is an axial structure, and in what others the appearance is due to +the adhesion of the base of the calyx to the carpels. + +The list which follows is not intended as a complete one, but it may +serve to show what plants are more particularly subject to this anomaly; +the * indicates unusual frequency of occurrence, the ! signifies that +the writer has himself seen instances in the plants named. Many of the +recorded cases of Synanthy are really cases of adhesion of the +inflorescence rather than of the flowers. + + Ranunculus Lingua. + bulbosus! + Aconitum Napellus. + Delphinium sp.! + Matthiola incana! + Arabis sagittata. + Silene sp. + Reseda odorata! + Vitis vinifera. + Citrus aurantium. + *Fuchsia var. hort.! + OEnothera sp. + Saxifraga sp. + Podalyria myrtillifolia. + Prunus Armeniaca. + spinosa. + Pyrus Malus. + Persica vulgaris. + Crataegus monogyna. + Robinia pseudacacia. + Gleditschia triacanthos. + Syringa persica. + Cornus sanguinea. + Viburnum sp. + *Lonicera sp. plur! + Centranthus ruber! + Valantia cruciata. + Centaurea moschata. + Jacea. + Zinnia elegans. + Zinnia revoluta. + Helianthus sp.! + Spilanthes oleracea. + Dahlia. + *Leontodon Taraxacum! + Senecio Doria. + Cichorium Intybus. + Lactuca sativa. + Anthemis retusa. + *Campanula medium! + persicifolia. + Azalea indica! + Vinca minor. + Atropa Belladonna. + *Solanum Lycopersicum! + *Petunia violacea! + Galeopsis ochroleuca. + Betonica alopecuros. + *Digitalis purpurea! + *Antirrhinum majus! + *Linaria purpurea! + *Pedicularis sylvatica! + *Calceolaria var. hort.! + Scrophularia nodosa. + Salpiglossis straminea. + Streptocarpus Rexii. + *Gesnera var. hort.! + AEschynanthus sp.! + Thyrsacanthus rutilans! + Anagallis collina. + *Primula veris! + Auricula. + *Primula acaulis, var. umbellata! + elatior? + * sinensis! + Aristolochia Clematitis. + Blitum sp. + Chenopodium sp. + Rumex sp. + Salix cinerea. + *Hyacinthus orientalis! + Lilium bulbiferum! + croceum, et sp. alix, pl. + Tulipa, sp. + Polygonatum anceps. + Fritillaria imperalis! + Agave americana. + Iris versicolor. + sambucina. + Crocus, sp. + Colchicum autumnale. + Narcissus incomparabilis! + Tazetta. + biflorus. + chrysanthus. + *Ophrys aranifera! + Calanthe vestita! + Oncidium bicolor. + ornithorhyncum. + &c. &c. + + In addition to the works before cited, additional information + on this subject may be gained from the following:--Jaeger, + 'Missbilld.,' p. 92. v. Schlechtend, 'Bot. Zeit.,' 1856, + _Robinia_. Weber, 'Verhandl. Nat. Hist. Vereins. Preuss. + Rheinl.,' 1849, p. 290, _Primula_. Hincks, 'Rep. Brit. Assoc. + Newcastle,' 1838, _Salpiglossis_. Clos, 'Mem. Acad. Toulouse,' + vol. vi, 1862, _Anagallis_. Wigand, 'Flora,' 1856, tab. 8, + _Pedicularis_. Henfrey, 'Botan. Gazette,' i, p. 280, _Reseda_. + P. Reinsch, 'Flora,' 1860, tab. 7, _Petasites_. Weber, + Verhandl. Nat. Hist. Vereins. f.d. Preuss. Rheinl. u. + Westphal.,' 1860, p. 332, tabs. 6 et 7, _Prunus_, _Persica_, + _Campanula_, _Taraxacum_, _Saxifraga_, _Silene_, _Hyacinthus_, + _&c._ Miquel, 'Linnaea,' xi, p. 423, _Colchicum_. Michel, + 'Traite du Citronnier,' tab. 6, _Citrus_. + +=Syncarpy.=--In the preceding section it has been shown that the +carpels, like other parts of the flower, are subject to be united +together. This union may either take place between the carpels of a +single flower or between the pistils of different flowers. In the latter +case the other floral whorls are generally more or less altered. Where, +however, the ovary is, as it is called, inferior, it may happen that the +pistils of different flowers may coalesce more or less without much +alteration in the other parts of the flower, as happens normally in many +_Caprifoliaceae_, _Rubiaceae_, &c. &c. In some of these cases it must be +remembered that the real structure of the apparent fruit is not made out +beyond dispute, the main points of controversy being as to what, if any, +share the dilated fruit-stalk or axis takes in the formation of such +organs. Again, it will be borne in mind that in some cases the so-called +fruit is made up of a number of flowers all fused together, as in the +Mulberry or the Pineapple, in which plants what is, in ordinary +language, called the fruit really consists of the whole mass of flowers +constituting the inflorescence fused together. Union of the fruits may +also in some cases take place between the carpels after the fall of the +other floral whorls, particularly when the outer layers of the pericarp +assume a succulent condition, so that under the general head of syncarpy +really different conditions are almost necessarily grouped together, +and, in seeking to investigate the causes of the phenomenon, the +particular circumstances of each individual case must be taken into +account. Syncarpy takes place in various degrees; sometimes only the +stalks are joined; at other times the whole extent of the fruit, as in +cherries, &c. This peculiarity did not escape the observant mind of +Shakespeare-- + + "A double cherry seeming parted. + But yet a union in partition, + Two lovely berries moulded on one stem." + 'Midsummer Night's Dream,' act iii, sc. 2. + +A similar union has been observed in peaches, gooseberries, gourds, +melons, and a great many other fruits. In the Barbarossa grape I have +frequently seen a fusion of two, three, four or more berries quite at +the end of the bunch, so that the clusters were terminated by a compound +grape. Seringe has remarked sometimes two, sometimes three, fruits of +_Ranunculus tripartitus_ soldered together. He has also seen three +melons similarly joined.[47] Turpin mentions having seen a complete +union between the three smooth and leathery pericarps which are +naturally separate and enclosed within the spiny cupule of the +chestnut.[48] Poiteau and Turpin have figured and described in their +treatise on fruit trees, under the name of Nefle de Correa, four or five +medlars, joined together and surmounted by all the persistent leaflets +of the calyces.[49] + +A very remarkable example of Syncarpy has been recorded by E. Koenig +in which nine strawberries were borne on one stem (_Fragaria +botryformis_),[50] and a similar malformation has been observed in the +Pineapple. + +When two fruits are united together they may be of about equal size, +while in other cases one of the two is much smaller than the other. This +was the case in two cucumbers given to me by Mr. James Salter. These +were united together along their whole length excepting at the very +tips; the upper one of the two was much larger than the lower, and +contained three cells, the lower fruit was one-celled by suppression. +Both fruits were curved, the curvature being evidently due to the more +rapid growth of the upper as compared with the lower one. + +[Illustration: FIG. 19.--Adhesion of two apples.] + +[Illustration: FIG. 20.--Section of united apples.] + +In many of these cases, where the fruits are united by their bases, the +summits become separated one from the other, so as to resemble the +letter V. Such divergence is of frequent occurrence where fruits are +united by their stalks, because, as growth goes on, the tendency must +necessarily be towards separation and divergence of the tips of the +fruit. + +In some cases of Syncarpy the fusion and interpenetration of the carpels +is carried to such an extent that it is very difficult to trace on the +outer surface the lines of union. The fruit in these cases resembles a +single one of much larger size than usual. Moquin mentions a double +apple in which the connection was so close that the fruit was not very +different in form from what is customary, and a similar thing happens +with the tomato. In the case of stone-fruits it sometimes happens, not +only that the outer portions are adherent, but that the stones are so +likewise. + +M. Roeper has observed two apples grafted together, one of which had +its stalk broken, and seemed evidently borne and nourished by the other +apple;[51] and a similar occurrence happens not infrequently in the +cucumber. Moquin has seen three united cherries having only a single +stalk jointed to the central fruit, the lateral cherries having each a +slight depression or cicatrix marking the situation of the suppressed +stalks. Schlotterbec has figured three apples presenting precisely +similar appearances.[52] + +Fusion of two or more nuts (_Corylus_) is not uncommon; I have seen as +many as five so united.[53] In these cases the fruits may be united +together in a ring or in linear series. + +In some _Leguminosae_, contrary to the general rule in the order, more +than one carpel is found; thus peas, French beans, and other similar +plants, are occasionally met with having two or more pods within the +same calyx, and in _Gleditschia triacanthos_ and _Caesalpinia digyna_ +this is so commonly the case as to be considered almost the normal +state. (De Cand. 'Mem. Leg.,' pl. 2, fig. 6; pl. 3, fig. 2.) At times +these carpels become fused together, and it becomes difficult, when the +traces of the flower have disappeared, to ascertain whether these +carpels were formed in one flower, or whether they were the result of +the fusion of several blossoms. I have seen an instance of this kind in +a plum in which there were two carpels in the same flower, the one being +partially fused to the other. The nature of such cases may usually be +determined by an inspection of the peduncle which shows no traces of +fusion. (See chapter on Multiplication.) + +When, however, the fruits are sessile, and they become grafted together, +the kind of syncarpy is difficult to distinguish. It, may, nevertheless, +be said as a general rule that the union brought about by the +approximation of two fruits, after the fall of the floral whorls, is +never so complete or so intimate as that determined by synanthy; and +also that in those cases where there are supernumerary carpels in the +flower, and those carpels become united together, they are rarely so +completely fused that their individuality is lost. + +An analogous phenomenon takes place not uncommonly in mosses, the spore +capsules of which become united together in various ways and degrees. +Schimper[54] cites the following species as subject to this +anomaly:--_Buxbaumia indusiata_, _Leskea sericea_, _Hypnum lutescens_, +_Anomodon alternatus_, _Clinacium dendroides_, _Bryum caespititium_, +_Brachythecium plumosum_, _Mnium serratum_, _Splachnum vasculosum_. It +has also been observed in _Trichostomum rigidulum_ and _Hypnum +triquetrum_. + + In addition to the authorities already mentioned, the reader + may consult Moquin-Tandon, 'El. Ter. Veg.,' p. 270. Turpin. + 'Mem. greffe. Ann. Sc. Nat.,' ser. i, t. xxiv, p. 334. De + Candolle, 'Organ. Veget.,' t. i. Duhamel, 'Phys. des Arbres,' + t. i, p. 304, tab. xiii, xiv. Weber. 'Verhandl. Nat. Hist. + Vereina f. d. Preuss. Rheinl. u. Westphal.,' 1860, p. 332, tab. + vi. et vii. + +=Synspermy, or Union of the Seeds.=--Seeds may be united together in +various degrees, either by their integuments,[55] or by their inner +parts. Such union of the seeds, however, is of rare occurrence. It takes +place normally, to a slight extent, in certain cultivated forms of +cotton, wherein the seeds are aggregated together into a reniform mass, +whence the term kidney cotton. Union of the parts of the embryo is +treated under another head (see Synophty). + +=Adhesion between the axes of different plants.=--Under this head may be +classed the union that takes place between the stems, branches, or roots +of different plants of the same species, and that which occurs between +individuals of different species; the first is not very different in its +nature from cohesion of the branches of the same plant (figs. 21, 22). +It finds its parallel, under natural circumstances, among the lower +cryptogams, in which it often happens that several individual plants, +originally distinct, become inseparably blended together into one mass. +In the gardening operations of inarching, and to some extent in budding, +this adhesion of axis to axis occurs, the union taking place the more +readily in proportion as the contact between the younger growing +portions of the two axes respectively is close. The huge size of some +trees has been, in some cases, attributed to the adnation of different +stems. This is said to be the case with the famous plane trees of +Bujukdere, near Constantinople, and in which nine trunks are more or +less united together.[56] + +[Illustration: FIG. 21.--Adhesion of two distinct stems of oak, or +possibly cohesion of branches of the same tree. 'Gard. Chron.,' 1846, p. +252.] + +A similar anastomosis may take place in the roots. Lindley cites a case +wherein two carrots, of the white Belgian and the red Surrey varieties +respectively, had grown so close to each other that each twisted half +round the other, so that they ultimately became soldered together; the +most singular thing with reference to this union was, that the red +carrot (fig. 23, _b_), with its small overgrown part above the +junction, took the colour and large dimensions of the white Belgian +(_d_), which, in like manner, with its larger head above the joining +(_a_), took the colour and small dimensions of the red one at and below +the union (_e d_). The respective qualities of the two roots were thus +transposed, while the upper portions or crowns were unaffected: the root +of one, naturally weak, became distended and enlarged by the abundant +matter poured into it by its new crown; and in like manner the root of +the other, naturally vigorous, was starved by insufficient food derived +from the new crown, and became diminutive and shrunken (see Synophty). + +[Illustration: FIG. 22.--Adhesion of the branches of two elms. 'Gard. +Chron.,' 1849, p. 421.] + +The explanation of the fact that the stumps of felled fir trees +occasionally continue to grow, and to deposit fresh zones of wood over +the stump, depends on similar facts. In _Abies pectinata_, says +Goeppert,[57] the roots of different individuals frequently unite; hence +if one be cut down, its stump may continue to live, being supplied with +nourishment from the adjacent trees to which it is adherent by means of +its roots. + +[Illustration: FIG. 23.--Adhesion of two roots of carrot. 'Gard. +Chron.,' 1851, p. 67.] + +A not uncommon malformation in mushrooms arises from the confluence of +their stalks (fig. 24), and when the union takes place by means of the +pilei, it sometimes happens, during growth, that the one fungus is +detached from its attachment to the ground, and is borne up with the +other, sometimes, even, being found in an inverted position on the top +of its fellow.[58] + +The garden operations of budding, grafting and inarching have already +been alluded to as furnishing illustrations of adhesion, but it may be +well to refer briefly to certain other interesting examples of adhesion +induced artificially; thus, the employment of the root as a stock, +"root-grafting," is now largely practised with some plants, as affording +a quicker means of propagation than by cuttings; and a still more +curious illustration may be cited in the fact that it has also been +found possible to graft a scion on the leaf in the orange.[59] + +[Illustration: FIG. 24.--Section through two adherent mushrooms, the +upper one inverted.] + +Mr. Darwin, in his work on the 'Variation of Animals and Plants,' vol. +i, p. 395, alludes to the two following remarkable cases of +fusion:--"The author of 'Des Jacinthes' (Amsterdam, 1768, p. 124) says +that bulbs of blue and red hyacinths may be cut in two, and that they +will grow together, and throw up a united stem (and this Mr. Darwin has +himself seen), with flowers of the two colours on the opposite sides. +But the remarkable point is, that flowers are sometimes produced with +the two colours blended together." In the second case related by Mr. +Trail, about sixty blue and white potatoes were cut in halves through +the eyes or buds, and the halves were then joined, the other buds being +destroyed. Union took place, and some of the united tubers produced +white, others blue, while some produced tubers partly white and partly +blue. + +=Adhesion of the axes of plants belonging to different species is a= +more singular occurrence than the former, and is of some interest as +connected with the operation of grafting. As a general rule +horticulturists are of opinion, and their opinion is borne out by facts, +that the operation of grafting, to be successful, must be practised on +plants of close botanical affinity. On the other hand, it is equally +true that some plants very closely allied cannot be propagated in this +manner. Contact between the younger growing tissues is essential to +successful grafting as practised by the gardener, and is probably quite +as necessary in those cases where the process takes place naturally. +Although there is little doubt but that some of the recorded instances +of natural or artificial grafting of plants of distant botanical +affinities are untrustworthy, yet the instances of adhesion between +widely different plants are too numerous and too well attested to allow +of doubt. Moreover, when parasitical plants are considered, such as the +Orobanches, the Cuscutas, and specially the mistleto (_Viscum_), which +may be found growing on plants of very varied botanical relationship, +the occurrence of occasional adhesion between plants of distant affinity +is not so much to be wondered at. Union between the haulms of wheat and +rye, and other grasses, has been recorded[60]. Moquin-Tandon[61] relates +a case wherein, by accident, a branch of a species of _Sophora_ passed +through the fork, made by two diverging branches of an elder +(_Sambucus_), growing in the Jardin des Plantes of Toulouse. The branch +of the _Sophora_ contracted a firm adhesion to the elder, and what is +remarkable is that, although the latter has much softer wood than the +former, yet the branch of the harder wooded tree was flattened, as if +subjected to great pressure[62]. It is possible that some of the cases +similar to those spoken of by Columella, Virgil[63], and other classical +writers, may have originated in the accidental admission of seeds into +the crevices of trees; in time the seeds grew, and as they did so, the +young plants contracted an adhesion to the supporting tree. Some of the +instances recorded by classical writers may be attributed to intentional +or accidental fallacy, as in the so-called "greffe des charlatans" of +more modern days. + +Adhesion of the roots of different species has been effected +artificially, as between the carrot and the beet root, while Dr. Maclean +succeeded in engrafting, on a red beet, a scion of the white Silesian +variety of the same species. In all these cases, even in the most +successful grafts, the amount of adhesion is very slight; the union in +no degree warrants the term fusion, it is little but simple contact of +similar tissues, while new growing matter is formed all round the cut +surfaces, so that the latter become gradually imbedded in the newly +formed matter. + +=Synophty or adhesion of the embryo.=--This often occurs partially in +the embryo plants of the common mistleto (_Viscum_), but is not of +common occurrence in other plants, even in such cases as the orange +(_Citrus_), the _Cycadeae_, _Coniferae_, &c., where there is frequently +more than one embryo in the seed. Alphonse De Candolle has described and +figured an instance of the kind in _Euphorbia helioscopia_, wherein two +embryo plants were completely grafted together throughout the whole +length of their axes, leaving merely the four cotyledons separate. A +similar adnation has been observed by the same botanist in _Lepidium +sativum_ and _Sinapis ramosa_, as well as in other plants.[64] I have +met with corresponding instances in _Antirrhinum majus_ and in _Crataegus +oxyacantha_, in the latter case complicated with the partial atrophy of +one of the four cotyledons. It is necessary to distinguish between such +cases and the fallacious appearances arising from a division of the +cotyledons. M. Morren has figured and described the union of two roots +of carrot (_Daucus_), which were also spirally twisted. He attributes +this union to the blending of two radicles, and applies the term +"rhizocollesy" to this union of the roots.[65] Mr. Thwaites cites a case +wherein two embryos were contained in one seed in a _Fuchsia_, and had +become adherent. What is still more remarkable, the two embryos were +different, a circumstance attributable to their hybrid origin, the seed +containing them being the result of the fertilisation of _Fuchsia +coccinea_ (quere _F. magellanica?_) by the pollen of _F. fulgens_. + +FOOTNOTES: + +[30] Wydler, 'Flora,' 1852, p. 737, tab. ix. + +[31] 'El. Ter. Veg.,' p. 254. + +[32] 'Bull. Soc. Bot. Fr.,' 1857, p. 451. + +[33] 'Bull. Acad. Belg.,' vol. xix, part ii, p. 335. + +[34] 'Bull. Soc. Bot. Fr.,' 1860, p. 25. + +[35] 'Bull. Soc. Bot. Fr.,' 1861, p. 147. + +[36] 'Bull. Acad. Belg.,' vol. xviii, part ii, p. 498. + +[37] See also Prillieux, 'Bull. Soc. Bot. Fr.,' 1861, p. 195. + +[38] 'Mem. Acad. Toulouse,' 5th Series, vol. iii. + +[39] Linnaea, vol. ii. p. 607. + +[40] 'Journal Roy. Hort. Soc.,' new ser., vol. i. 1866, p. 200. + +[41] 'Bull. Soc. Bot. Fr.,' 1861, p. 159. + +[42] Ibid., 1859, p. 467. + +[43] 'Flora,' 1858, p. 65, tab. ii. + +[44] C. Morren. 'Bull. Acad. Belg.,' vol. xv (Fuchsia, p. 89); vol. +xviii, p. 591. (Lobelia, p. 142); vol. xix, p. 352; vol. xx, p. 4. + +[45] 'Bull. Soc. Bot. Fr.,' vol. vii, p. 625. + +[46] Cramer, 'Bildungsabweichungen,' p. 56, tab. vii, fig. 10, figures a +case wherein the two central flowers of the capitulum of _Centaurea +Jacea_ were united together. + +[47] 'Bull. Bot.' tab. iii, figs. 4-6. + +[48] 'Mem. greffe Ann. Science Nat.,' ser. i, t. xxiv, p. 334. + +[49] "Mespilus portentosa." Poit. et Turp., 'Pomol. Franc.,' liv, xxxi, +p. 202, pl. 202. + +[50] Duchesne, 'Hist. Nat. Frais.,' p. 79. + +[51] De Cand., 'Phys. Veget.,' tom. ii, p. 781. + +[52] Sched. de monstr. plant. 'Act. Helv.,' tab. i, fig. 8. + +[53] 'Mem. greffe,' loc. cit., tab. xxiv, p. 334. + +[54] 'Bull. Soc. Bot. Franc.,' 8, pp. 73 and 351, tab. ii; and Roese. +'Bot. Zeit.,' x, p. 410. + +[55] _Nymphaea lutea_, _AEsculus Hippocastanum_, &c. See Moquin, 'El. +Ter. Veg.,' p. 277. + +[56] C. Martins, 'Promenade Botanique,' p. 8. + +[57] 'Ann. Sc. Nat.,' t. xix, 1843, p. 141, tab. iv. + +[58] 'Ann. Nat. Hist.,' ser. 2, vol. ix, tab. xvi. 'Phytologist,' 1857. +p. 352, &c. + +[59] Quoted from the 'Revue Hortic.' in 'Gard. Chron.,' 1866, p. 386. + +[60] Senebier, 'Phys Veget.,' t. iv, p. 426. The same author also cites +Romer as having found two plants of _Ranunculus_, from the stem of which +emerged a daisy. As it is not an uncommon practice to stick a daisy on a +buttercup, it is to be hoped no hoax was played off on M. Romer. + +[61] 'El. Ter. Veg.,' p. 289. + +[62] An instance of this kind is cited in Dr. Robson's memoir of the +late Charles Waterton, from which it appears that two trees, a spruce +fir and an elm, were originally planted side by side, and had been +annually twisted round each other, so that they had in places grown one +into the other, with the result of stunting the growth of both trees, +thus illustrating, according to the opinion of the eccentric naturalist +above cited, the incongruous union of Church and State! + +[63] See Daubeny, 'Lectures on Roman Husbandry,' p. 156. + +[64] A. P. De Candolle, 'Organ Veget.,' t. ii, p. 72, tab. liv, fig. 1. + +[65] 'Bull. Acad. Belg.,' t. xx, part i, 1852, p. 43. + + + + +PART II. + +INDEPENDENCE OR SEPARATION OF ORGANS. + + +Under this head are included all those instances wherein organs usually +entire, or more or less united, are, or appear to be, split or +disunited. It thus includes such cases as the division of an ordinarily +entire leaf into a lobed or partite one, as well as those characterised +by the separation of organs usually joined together. Union, as has been +stated in a previous chapter, is the result either of persistent +integrity or of a junction of originally separate organs, after their +formation; so in like manner, the separation or disjunction of parts may +arise from the absence of that process of union which is habitual in +some cases, or from an actual _bona fide_ separation of parts originally +united together. In the former case, the isolation of parts arises from +arrest of development, while in the latter it is due rather to luxuriant +growth. A knowledge, as well of the ordinary as of the unusual course, +of development in any particular flower is thus required in order to +ascertain with accuracy the true nature of the separation of parts. The +late Professor Morren[66] proposed the general term Monosy ([Greek: +monosis]) for all these cases of abnormal isolation, subdividing the +group into two, as follows--1, Adesmy ([Greek: a-desmos]), including +those cases where the separation is congenital; and 2, Dialysis ([Greek: +dialyo]), comprising those instances where the isolation is truly a +result of the separation of parts previously joined together. Adesmy, +moreover, was by the Belgian savant said to be homologous when it +occurred between members of the same whorl, _e.g._ between the sepals of +an ordinary monosepalous calyx, or heterologous when the separation took +place between members of different whorls, as when the calyx is detached +from the ovary, &c. The former case would thus be the converse of +cohesion, the latter of adhesion. + +To the adoption of these words there is this great objection, that we +can but rarely, in the present state of our knowledge, tell in which +group any particular illustration should be placed. + +The terms adopted in the present work are, for the most part, not +necessarily intended to convey any idea as to the organogenetic history +of the parts affected. Where a single organ, that is usually entire, +becomes divided the term Fission is used; in cases where parts of the +same whorl become isolated, the word Dialysis is employed, and in the +same sense in which it is generally used by descriptive botanists, and +where the various whorls become detached one from the other, the +occurrence is distinguished by the application of the term Solution. + +FOOTNOTES: + +[66] 'Bull. Acad. Belg.,' t. xix, part iii, 1852, p. 315. + + + + +CHAPTER I. + +FISSION. + + +When an organ becomes divided it receives at the hands of descriptive +botanists the appellations cleft, partite, or sect, according to the +depth of the division; hence in considering the teratological instances +of this nature, the term fission has suggested itself as an appropriate +one to be applied to the subdivision of an habitually entire or +undivided organ. It thus corresponds pretty nearly in its application +with the term Chorisis or "dedoublement," or with the "disjonctions qui +divisent les organes" of Moquin-Tandon.[67] It is usually, but not +always, a concomitant with hypertrophy, and dependent on luxuriance of +growth. + +It must be understood therefore that the term, as generally applied, +does not so much indicate the cleavage of a persistent organ, as it does +the formation and development of two or more growing points instead of +one, whence results a branching or forking (di-tri-chotomy) of the +affected organ. In some instances it seems rather to be due to the +relative deficiency of cellular, as contrasted with fibro-vascular +tissue. + +=Fission of axile organs.=--This condition is scarcely to be +distinguished from multiplication of the axile organs (which see). A +little attention, however, will generally show whether the unusual +number of branches is a consequence of the development of a large number +of distinct shoots, as happens, for instance, when a tree is pollarded, +or of a division of one. M. Fournier[68] gives as an illustration the +case of a specimen of _Ruscus aculeatus_ in which there occurred a +division of the foliaceous branches into two segments, reaching as far +as the insertion of the flower, but no further. He also mentions lateral +cleavage effected by a notching of the margin, the notch being anterior +to the flowers and always directed towards their insertion. In the +allied genus _Danae_, Webb, 'Phyt. Canar.,' p. 320, describes the +fascicles of flowers as in "crenulis brevibus ad marginem ramulorum +dispositis." Sometimes, on the other hand, _Danae_ has a fascicle of +flowers inserted on the middle of the upper surface, as in _Ruscus_. +Wigand mentions an instance in _Digitalis lutea_, where the upper part +of the stem was divided into six or seven racemes; possibly this was a +case of fasciation, but such a division of the inflorescence is by no +means uncommon in the spicate species of _Veronica_. I have also seen +it in _Plantago lanceolata_, _Reseda luteola_, _Campanula medium_, +_Epacris impressa_, and a bifurcation of the axis of the spikelet within +the outer glumes in _Lolium perenne_[69] and _Anthoxanthum odoratum_. In +the Kew Museum is preserved a cone of _Abies excelsa_,[70] dividing into +two divisions, each bearing bracts and scales. A similar thing +frequently occurs in the male catkins of _Cedrus Libani_ (fig. 25). + +[Illustration: FIG. 25.--Bifurcated male inflorescence, _Cedrus +Libani_.] + +This subdivision of axial organs is not unfrequently the result of some +injury or mutilation, thus Duval Jouve alludes to the frequency with +which branched stems are produced in the various species of _Equisetum_, +as a consequence of injuries to the main stem, but this is rather to be +considered as a multiplication of parts than as a subdivision of one. + +[Illustration: FIG. 26.--Bifurcated leaf of _Lamium album_, &c.] + +=Fission of foliar organs.=--Many leaves exhibit constantly the process +of fission, such as the _Salisburia adiantifolia_, and which is due +perhaps as much to the absence or relatively small proportion of +cellular as compared with vascular tissue, as to absolute fission. In +the same way we have laciniated leaves of the Persian lilac, _Syringa +persica_, and Moquin mentions instances in a species of _Mercurialis_ +in which the leaves were deeply slashed. In _Chenopodium Quinoa_ the +leaves were so numerous and the clefts so deep, that the species was +hardly recognisable, while on a branch of _Rhus Cotinus_ observed by De +Candolle the lobes were so narrow and so fine as to give the plant the +aspect of an _Umbellifer_. Wigand ('Flora,' 1856, p. 706) speaks of the +leaves of _Dipsacus fullonum_ with bi-partite leaves; Moquin mentions +the occurrence of a leaf of an oleander bi-lobed at the summit, so as to +give the appearance of a fusion of two leaves. Steinheil has recorded an +instance in _Scabiosa atropurpurea_ in which one of the stem leaves +presented the following peculiarities. It was simple below, but divided +above into two equal lobes, provided each with a median nerve.[71] +Steinheil has also recorded a _Cerastium_ in which one of the leaves was +provided with two midribs; above this leaf was a group of ternate +leaves. I have seen similar instances in the common Elm, _Ulmus +campestris_, and also in the common nettle, _Urtica dioica_, the leaves +of which latter thus resembled those of _Urtica biloba_, which are +habitually bilobed at the summit. M. Clos[72] mentions an instance where +the terminal leaf and first bract of _Orchis sambucina_ were divided +into two segments. The same author also mentions the leaves of +_Anemiopsis californica_, which were divided in their upper halves each +into two lobes--also leaves of a lentil springing from a fasciated stem +and completely divided into two segments, but with only a single bud in +the axil. The axillary branches in like manner showed traces of +cleavage. Fig. 26 represents a case of this kind in _Lamium album_, +conjoined with suppression of the flowers on one side of the stem. I +have also in my herbarium a leaf of _Arum maculatum_, with a stalk +single at the base, but dividing into two separate stalks, each bearing +a hastate lamina, the form of which is so perfect that were it not from +the venation of the sheath it would be considered that there was here a +union of two leaves rather than a bifurcation of one. A garden +Pelargonium presented the same appearance. + +[Illustration: FIG. 27.--Bifurcated leaf of _Pelargonium_.] + +Fern fronds are particularly liable to this kind of subdivision, and +they exhibit it in almost every degree, from a simple bifurcation of the +frond to the formation of large tufts of small lobes all formed on the +same plan by the repeated forking of the pinnules. These may be +considered as cases of hypertrophy. + +Moquin-Tandon, at a meeting of the Botanical Society of France (April +3rd, 1858) exhibited a leaf of _Cerasus Lauro-Cerasus_ divided in such a +manner as to resemble a leaf of _Citrus_ or of _Phyllarthron_. In this +case, therefore, the disunion must have taken place laterally, and not +from apex towards base, as is most common. The leaves of the common +horse-radish, _Cochlearia Armoracia_, are very subject to this pinnated +subdivision of the margin, and numerous other illustrations might be +given. + +[Illustration: FIG. 28.--Bifurcated frond, _Scolopendrium vulgare_.] + +A. Braun describes a singular case in a leaf of _Irina glabra_ +wherein the blade of the leaf on one side was deeply and irregularly +laciniated, the other side remaining entire. (Verhandl., d. 35, +Naturforscherversammlung, tab. 3.) Laciniate varieties of plants are of +frequent occurrence in gardens where they are often cultivated for their +beauty or singularity; thus, there are laciniated alders, fern-leaved +beeches and limes, oak-leaved laburnums, &c. A list of several of these +is subjoined. A similar fission takes place constantly in the cotyledons +of some plants, sometimes, as in _Coniferae_, to such an extent as to +give an appearance as if there were several cotyledons.[73] + +It is not always easy to recognise, at a first glance, whether the +division be the result of disunion or of an incomplete union of two +leaves, but we may be guided by the number of leaves in the cycle or the +whorl. The number is complete in cases of partial disjunction, while in +cases of fusion it is incomplete. Again, in instances of disjunction, +there is only one point of origin, but, when two leaves are grafted +together, two such points may generally be detected at the base of the +leaf, or a transverse section of the leaf-stalk will show indications of +fusion. The number and position of the midribs will also serve as a +guide, as in cases of fusion there are generally two or more midribs, +according to the number of fused leaves; but as Moquin well remarks, +this latter character cannot be always depended upon, for the median +nerve may divide without any corresponding separation of the cellular +portions of the leaf. The author just quoted cites examples of this kind +in _Cardamine pratensis_, _Hedera Helix_, _Plantago major_, _Geranium +nodosum_. + +The following list of plants commonly producing leaves that are cleft or +divided, to a greater extent than is usual in the species, is mainly +taken from one given by Schlechtendal, 'Bot. Zeit.,' 1844, p. 441, with +additions from other sources. The ! indicates that the author has +himself met with the deviation in question. Many are cultivated as +garden varieties under the names here given. + + Trollius europaeus dissectus. + Chelidonium majus laciniatum! + Glaucium luteum. + Brassica oleracea! + Tilia parvifolia laciniata. + asplenifolia! + Acer platanoides laciniatum. + crispum. + AEsculus Hippocastanum incisum! + asplenifolium. + Vitis vinifera apiifolia! + laciniosa. + Ilex Aquifolium! + Rhus Toxicodendron quercifolium. + Cotinus. + Ervum Lens. + Cytisus Laburnum quercifolium! + incisum. + Rubus fraticosus laciniatus! + Pyrcis communis. + Cerasus Lauro-cerasus. + Apium graveolens! + Pimpinella magna. + Saxifraga. + Crataegus Oxyacantha laciniata. + quercifolia! + Ribes nigrum. + Sambucus nigra laciniata! + racemosa laciniata. + Dipsacus fullonum. + Scabiosa atropurpurea! + Symphoricarpus racemosus. + Helianthus sp.! + Lonicera Periclymenum quercifolia! + Syringa persica laciniata! + Syringa vulgaris! + Nerium Oleander! + Lamium purpureum. + album! + Salvia officinalis. + Solanum Dulcamara! + Fraxinus excelsior crispa. + Veronica austriaca. + Polemonium caeruleum. + Juglans regia laciniata! + heterophylla. + filicifolia. + Anemiopsis californica. + Chenopodium Quinoa. + Ulmus americana incisa. + Fagus sylvatica heterophylla! + laciniata! + aspleniifolia! + incisa. + salicifolia! + Mercurialis perennis. + Urtica dioica. + Quercus Cerris laciniata! + pubescens filicina. + Betula populifolia laciniata. + alba dalecarlica. + Alnus incana laciniata! + glutinosa laciniata! + quercifolia. + oxyacanthifolia. + Corylus Avellana heterophylla! + laciniata! + urticifolia. + Carpinus Betulus incisa! + quercifolia. + heterophylla. + Castanea vesca heterophylla. + quercifolia. + incisa. + Populus alba acerifolia. + palmata. + quercifolia. + balsamifera. + Orchis sambucina. + Arum maculatum. + Filices sp. pl. + + See also Schlechtendal, 'Bot. Zeit.,' tom. xiii, p. 823. A. + Braun, loc. supra citat. For Ferns too numerous for insertion, + see Moore, 'Nature-Printed Ferns,' 8vo ed., 2 vols. Clos, 'Mem. + Acad. Toulouse,' 1862, p. 51. + +=Fission of the petals, &c.=--The floral leaves are subject to a similar +process of cleavage to that which has just been mentioned as taking +place in the leaves. This, indeed, occurs very often as a normal +occurrence as in the petals of mignonette (_Reseda_), or those of +_Alsine media_ and many other plants. Here, however, we have only to +allude to those instances in which the cleavage occurs in flowers whose +sepals or petals are usually entire. Under this category Moquin mentions +a petal of _Brassica oleracea_ completely split into two. Linne in his +'Flora Lapponica' (pp. 145 and 164) mentions quadrifid petals of +_Lychnis dioica_, and much divided petals of _Rubus arcticus_. Among +other plants subject to this division of sepals or petals may be +mentioned as having come within the writer's personal observation, +_Ranunculus Lingua_, _R. acris_, _Papaver somniferum_, and others of +this genus, _Saponaria sp._, _Dianthus_, _Narcissus_, &c. + +In some of the garden varieties of _Cyclamen_ the corolla looks at first +sight as if double, and the plan of the flower is oblong or elliptical, +instead of circular. In these flowers each lobe of the corolla is +divided almost to the base into two lobes, so that there appear to be +ten lobes to the corolla instead of five, as usual. The stamens are +normal in form and number in these flowers. + +In the paroquet tulips of gardeners the segments of the perianth are +deeply and irregularly gashed, the segments occasionally becoming rolled +up and their margins coherent so as to form little tubular spurs. I have +also noticed the segments of the perianth in _Crocus_ and _Colchicum_ +deeply cleft, so much so sometimes, as to equal in this particular the +stigmas. In the flowers of a species of _Oncidium_, communicated to me +by Mr. Currey, the lip was divided into three segments perfectly +distinct one from the other, but confluent with the column; the two side +pieces had callosities at the upper edge close to the base, the central +piece had a similar wartlike process in its centre. In these flowers the +ovary, the stigma, and the anther were all in a rudimentary condition. +Some verbenas raised by Mr. Wills offer a curious illustration of this +condition. It will be remembered that some of the lobes or petals of a +verbena are normally divided at the base to a slight degree, but in the +flowers in question this is carried to such an extent that the enlarged +lobes are pushed into the centre of the flower and simulate, at a first +glance, a distinct and separate organ, though in reality it is but an +enlargement of what occurs normally.[74] + +[Illustration: FIG. 29.--Flower of _Oncidium sp._ seen from the back. +The lip is divided into three unequal segments.] + +Moquin mentions having seen the stamens of _Matthiola incana_ and +_Silene conica_ completely divided, each section bearing half an anther, +exactly as happens in _Polygalaceae_. In tulips and lilies the same +author mentions division of the anther only, the filament remaining +entire, as happens naturally in many species of _Vaccinium_. + +A division of the individual carpels occurs very frequently when those +organs become more or less leafy, as in _Trifolium repens_, and other +plants to be hereafter mentioned. + +The instances given in this chapter have all been cases wherein the +division or the accessory growth has taken place in one plane only and +that plane the same as that of the affected organ, but there are other +examples, probably equally due to fissiparous division, where the new +growth is either parallel to, or even at angle with the primary organ. +Of such nature are some of those instances wherein two leaves appear to +be placed back to back. These partake of the nature of excrescences or +of exaggerated developments, and hence will be more fully treated of +under the head of hypertrophy. It must be remembered that in some of +these cases the fission may be a resumption of characters proper to the +species under natural conditions, but lost by cultivation or otherwise. +Thus, Mr. Buckman accounts for "finger-and-toe" in root-crops on the +principle of reversion to the wild form. + +FOOTNOTES: + +[67] Loc. cit., p. 295. + +[68] 'Bull. Soc. Bot. France,' 1857, p. 758. + +[69] Masters, 'Jourl. Linn. Soc.,' vol. vii, p. 121. + +[70] Cramer, 'Bildungsabweichungen,' p. 4, tab. vi, fig. 4, figures a +case of the same kind in _Pinus Cembra_. + +[71] 'Ann. des Science Nat.,' 2nd series, t. iv, p. 147, tab. v, figs. 3 +and 4. + +[72] 'Mem. Acad. Scien. Toulouse,' 5th series, vol. iii. + +[73] Duchartre, 'Ann. Sc. Nat.,' 3rd series, 1848, vol. x, p. 207. + +[74] Masters, 'Rep. Bot. Congress,' London, 1866, p. 136, tab. 7, f. 15, +16. + + + + +CHAPTER II. + +DIALYSIS. + + +This term is here made use of in the same sense as in descriptive +botany, to indicate the isolation of parts of the same whorl; it is thus +the opposite of cohesion. Morren, as has been previously stated, +employed the word in a different sense, while Moquin-Tandon[75] included +cases of this description under the category of "Disjonctions qui +isolent les organes." + +Dialysis, as here understood, may be the result of an arrest of +development, in consequence of which parts that under ordinary +circumstances would become fused, do not do so; or, on the other hand, +it may be the result of an actual separation between parts primitively +undivided. As it is not possible in every case to distinguish between +the effects of these two diverse causes, no attempt is here made to do +so. + +=Dialysis of the margins of individual foliar organs.=--In cases where +the leaf or leaf-like organ is ordinarily tubular or horn-like in form, +owing to the cohesion of its edges, it may happen either from lack of +union or from actual separation of the previously united edges, that the +tubular shape is replaced by the ordinary flattened expansion. Thus, in +_Eranthis hyemalis_, wherein the petals (nectaries) are tubular and the +sepals flat, I have met with numerous instances of transition from the +one form to the other, as shown in fig. 9, p. 24. + +It is, however, in the carpels that this separation occurs most +frequently. When these organs appear under the guise of leaves, as they +often do, their margins are disunited, so that the carpel becomes flat +or open. This happens in the strawberry (_Fragaria_), the columbine +(_Aquilegia_), in _Trifolium repens_, _Ranunculus Ficaria_, &c.[76] + +=Dialysis of the parts of the same whorl:--calyx.=--The separation of an +ordinarily coherent series into its constituent parts is necessarily of +more common occurrence than the foregoing. As here understood, it is the +precise converse of cohesion, and it may be represented diagrammatically +by a dotted line above the letters denoting the sepals, petals, &c. When +this change happens in the calyx we have the gamosepalous condition +replaced by the polysepalous one, as thus represented: + + ............. + S S S S S +instead of + _____________ + S S S S S + +as in a calyx of five coherent sepals. + +Detachment of this kind occurs not unfrequently, as in _Primula +vulgaris_, _Trifolium repens_, &c. In _Rosaceae_ and _Pomaceae_ this +separation of the calyx is of the more moment, as it has reference to +the structure of the inferior ovary, as will be more fully mentioned +hereafter. Here, however, a case recorded by M. J. E. Planchon may be +alluded to[77] wherein a quince fruit (_Cydonia_) was surmounted by five +leaves, the surface of the pome being marked by as many prominences, +which apparently corresponded to the five stalks of the calycine leaves. +In this specimen, then, the inferior position of the ovary appeared to +be not so much due to an expansion of the fruit stalk, as to the fusion +of the hypertrophied stalks of the sepals. Some of the malformations +among Cucurbits point to a similar structure. It is probable that in +many of these cases the so-called inferior ovary is partly axial partly +foliar, _i.e._, sepaline, and partly carpellary in its nature. + +Dialysis of the sepals in calyces that are usually gamosepalous has been +most frequently observed in _Rosaceae_, _Pomaceae_, _Umbelliferae_, less +commonly in _Leguminosae_, also in the following genera:--_Primula_, +_Symphytum_, _Gentiana_, _Campanula_, &c. + +[Illustration: FIG. 30.--Dialysis of the sepals and petals in _Correa_.] + +=Dialysis of the corolla= is likewise of frequent occurrence, either +partially or to such an extent as to render the corolla truly +polypetalous. Among _Labiatae_ the upper lip of the corolla may be often +met with partially cleft, as it is constantly in _Phlomis biloba_, or +more markedly among the _Lobeliaceae_. + +In the _Compositae_, a similar separation of the petals is not +infrequent, thus showing frequent transitional stages between the +labiatifloral and tubulifloral divisions respectively. The ligulate +corollas also may often be found in Chrysanthemums, Dahlias, &c., more +or less deeply divided into their component parts. + +A more complete separation occurs not unfrequently in _Campanula_, +_Rhododendron_, _Phlox_, _&c._ Figs. 30 and 31 illustrate dialysis of +the corolla; the first in _Correa_, the second in _Campanula_. + +[Illustration: FIG. 31.--Dialysis of the corolla in _Campanula sp._, +after De Candolle.] + +In the last-named genus, _C. rotundifolia_ has been found with +polypetalous flowers in a wild state in the mountains of Canton +Neufchatel, Switzerland, and gave rise to the creation of a new genus. +This form is now introduced into gardens. + +It must be remembered that in some genera, where this separation of the +petals has been met with, there are species in which a similar isolation +occurs normally, as in _Rhododendron_. _R. linearilobum_, a Japanese +species, offers a good illustration of this. + +The following list contains the names of the genera in which this +separation of the petals of an ordinarily gamopetalous flower takes +place most frequently. + + Correa. + Campanula! sp. pl. + Polemonium. + Phlox! + Coboea! + Rhododendron! + Erica! + Rhodora. + Azalea! + Compositae! sp. pl. + Lonicera! + Convolvulus! + Pharbitis. + Antirrhinum! + Verbascum! + Mimulus. + Digitalis! + Orobanche. + Solanum. + Nicotiana. + Gentiana! + Anagallis. + Primula! + Lamium! + Convallaria! + Lilium! + Colchicum! + &c. &c. + +This list does not include those very numerous cases in which this +change is associated with more or less complete frondescence or leafy +condition of the petals. + +=Dialysis of the stamens.=--A similar isolation of the stamens occurs +occasionally; for instance, when Mallows (_Malvaceae_) become double, one +of the first stages of the process is often the disjunction of the +stamens, and a similar dissociation occurs in _Leguminosae_ and +_Compositae_, as in _Tragopogon_, as related by Kirschleger, in +_Hypochaeris_ by Wigand, and in _Coreopsis_ by Schlechtendal. + +=Dialysis of the carpels.=--In the case of the carpels this disunion is +more frequent than in the stamens. M. Seringe[78] figures carpels of +_Diplotaxis tenuifolia_ more or less completely separated one from the +other; indeed, this separation is very common amongst _Cruciferae_ and +_Umbelliferae_. + +Generally speaking, the disunion is complicated with frondescence--but +not always so. I have, in my herbarium, specimens of _Convallaria +majalis_, _Commelyna sp._, and of _Lilium auratum_, in all of which the +three carpels are completely disjoined, and present three styles, three +stigmas, &c., without any other change. Engelmann[79] speaks of three +classes of this malformation. 1st, that in which the carpels separate +one from the other without opening, as in the lily just alluded to; 2nd, +that in which the ovary remains closed, but loses its internal +partitions, as in a case mentioned by Moquin in _Stachys sylvatica_, in +which, owing to imperfect disjunction, the two bi-lobed carpels were +changed into a nearly one-celled capsule;[80] and 3rd, those cases in +which the carpels are open and foliaceous. + +[Illustration: FIG. 32.--Anomalous form of orange.] + +Disjunction is more frequent in dry fruits than in fleshy ones. In the +latter instance it happens at an early stage of existence, and the +pericarp becomes more or less leafy, losing its faculty of becoming +fleshy, as in _Prunus Cerasus_ and _Amygdalus persica_; nevertheless, +fleshy fruits sometimes become disunited. I have seen a case similar to +that mentioned by M. Alphonse de Candolle in _Solanum esculentum_, in +which the pericarp became ruptured, and the placentas protruded. A like +occurrence has also been observed in a species of _Melastoma_.[81] This +is analogous to what happens in _Caulophyllum_ and _Slateria_. +Disjunction of the carpels is not rare in oranges. Sometimes this takes +place regularly, at other times irregularly; occasionally in such a +manner as to give the appearance of a hand and fingers to the fruit. Of +one of these, Ferrari,[82] in the curious volume below cited, speaks +thus: "Arbor profusissima, quia dat utraque manu; imo quia vere manus +dat in poma conversis; utque magis munifica sit poma ipsa convertit in +manus." + +M. Duchartre[83] mentions a semi-double flower of orange with eight to +ten distinct carpels in a whorl, and occasionally several whorls one +above another. De Candolle[84] considers the rind of the orange as a +production from the receptacle, and this view is confirmed by the +specimens of Duchartre, in which the carpels were quite naked or had a +common envelope truncated, and open above to allow of the passage of the +styles and stigmas. + +[Illustration: FIG. 33.--Orange. Showing disjunction of carpels, after +Maout.] + +[Illustration: FIG. 34.--Section of orange shown in fig. 33 after +Maout.] + +It frequently happens in conjunction with this separation of the carpels +one from the other, that a lack of union manifests itself between the +margins of the individual carpels themselves. Very numerous cases of +this kind have been recorded, and the double tulips of gardens may be +referred to as showing this condition very frequently. In connection +with this detachment of the carpels, a change in the mode of +placentation is often to be observed, or two or more kinds may be seen +in the same pistil, as in double-flowered saponarias, many Crucifers, +&c., as alluded to under the head of displacements of the placenta. + +FOOTNOTES: + +[75] _Loc. cit._, p. 298. + +[76] Masters in Seemann's 'Journal of Botany,' 1867, p. 158. + +[77] Bull. Soc. Bot. France,' t. xiii, 1866, p. 234. + +[78] 'Bull. Bot.,' pl. i, figs. 8-12. + +[79] 'De Anthol.,' p. 37. + +[80] Moquin, loc. cit., p. 305. + +[81] 'Neue Denkschr. der Allg. Schweiz. Gesell.,' band v, pl. ii. p. 5. + +[82] 'Hesperides,' auctore Ferrario. Rome, 1646, fig. 415, pp. 213 and +215. See also Michel, 'Traite du Citronnier.' + +[83] 'Ann. des Science Nat.,' 3rd series, 1844, vol. i, p. 294. + +[84] 'Org. Veget.,' vol. ii., p. 41. + + + + +CHAPTER III. + +SOLUTION. + + +The isolation or separation of different whorls that are ordinarily +adherent together is by no means of rare occurrence. Were it not that +the isolation is often congenital, the word detachment would be an +expressive one to apply to these cases, but as the change in question +occurs quite as often from a want of union, an arrest or stasis of +development, as from a _bona fide_ separation, the word solution seems +to be, on the whole, the best. It corresponds in application to the word +_liber_ (_calyx liber_, &c.), in general use by descriptive botanists. +As here employed, the term nearly corresponds with the "adesmie +heterologue" of Morren. Moquin Tandon does not make any special +subdivision for the class of cases here grouped together, but places +them all under "Disjonctions qui isolent les organes." It seems, +however, desirable to have a separate word to express the converse +condition of adhesion, and for this purpose the term solution, as above +stated, is here employed. Diagrammatically, the condition may be +expressed by placing a dotted line at the side of the letters thus: + + : s s s s s : + : c c c c c : + +would indicate the disjunction of the sepals from the carpels (c), in +contradistinction to adhesion, which may be represented by the unbroken +line thus: + + | s s s s s | + | c c c c c | + +=Solution of the calyx from the ovary.=--Of all the instances of +adhesion which take place under ordinary circumstances, that between the +calyx and the ovary is perhaps the most common. The _calyx adhaerens_ or +_superus_ is a structural characteristic to which all botanists attach +considerable importance; so that when exceptional cases occur in which +the calyx becomes detached from the ovary, becomes, that is, _inferus_ +or _liber_, a proportionate degree of interest attaches to the +irregularity. It is not within the scope of the present work to inquire +whether this detachment be real or merely apparent, arising from a want +of union between parts ordinarily united together. This point must be +left to the organogenists to decide in each particular case. So also the +question as to what share, if any, the expanded and dilated flower-stalk +may take in what are usually called inferior ovaries, can be here only +incidentally touched upon. + +Among _Rosaceae_, the change in question is very common, especially in +conjunction with an elongation of the axis of the flower (apostasis) and +with prolification, though it is by no means always co-existent with +these malformations. When this alteration in the apparent relative +position of calyx and carpels occurs in roses (_Rosa_) the appearances +are generally such as to indicate that the "hip" of the rose is a +dilatation of the peduncle, continuous above with the coherent bases of +the sepals; this inference seems also to be borne out by what happens in +the _Pomaceae_. In some cases in this sub-order, the calyx becomes +detached from the carpels, so that the latter organs become more or less +"superior," and distinct one from the other. This happens constantly in +the double-flowered thorn, _Crataegus Oxyacantha_, in some blossoms of +which the hollowed end of the peduncle still invests the base of the +carpels, leaving the upper portions detached. In apples flowers are +occasionally met with of greater size than usual and on longer stalks, +so that the whole looks more like a rose than an apple blossom. In these +cases it will usually be found that the calyx consists of distinct +sepals, without a trace of the ordinary swelling beneath the flower. The +petals are often more numerous than usual; the stamens variously +changed, and the carpels sometimes absent; at other times, as in the +instance figured in the adjacent woodcuts, figs. 36, 37, consisting of +separate, superior ovaries, sometimes destitute of ovules, or, at other +times, having two of these bodies.[85] + +[Illustration: FIG. 35.--Proliferous Rose. Showing an absence of the +usual dilatation of the flower-stalk, and other changes.] + +This condition accords precisely with the account of the development of +the flowers in _Pomaceae_ as given by Payer, Caspary, and others, so that +the flowers above described would owe their deficiency of the swollen +receptacle to an arrest of development. M. Germain de Saint Pierre, +among other malformations of the rose, presented to the Botanical +Society of France in 1854[86] two specimens which are of special +interest as relating to this contested point. In the one, the swollen +portion beneath the flower was surmounted by five perfect leaves, as, +indeed, is not infrequent in such malformations; here, then, the calyx +could have had little or no share in the production of the swelling in +question. In the other, the swollen portion was actually above the +insertion of the sepals here represented by five perfect leaves. + +[Illustration: FIG. 36.--Section through Apple blossom, showing +detachment of calyx from ovaries, absence of dilated flower-stalk, &c.] + +[Illustration: FIG. 37.--Calyx detached from carpels in Apple.] + +On the other hand, M. Planchon's specimen of the Quince before alluded +to, not to mention other instances, tends to show that the bases of the +sepals do sometimes enter into the composition of the pome. And, indeed, +in many of these cases it would be impossible to say where the axial or +receptacular portion ended, and the foliar portion began. As both from +normal organogeny as well as from unusual conformation contradictory +inferences may be drawn, it would obviously be unsafe to attempt the +explanation of the so-called calyx-tube in general from any particular +instances; so far as _Rosaceae_ are concerned, there is so much variation +in the relative position of calyx and carpels under ordinary +circumstances, that it is no matter for surprise that similar +diversities should exist in teratological cases. A similar remark will +apply to _Saxifragaceae_, _Cucurbitaceae_, _Myrtaceae_, _Bruniaceae_, +_Rubiaceae_, and other families of like conformation. + +[Illustration: FIG. 38.--Flower of _OEnanthe crocata_, in which the +five sepals were completely detached from the ovaries, here three in +number and destitute of stylopods.] + +In _Umbelliferae_, a detachment of the calyx from the ovaries frequently +occurs, sometimes without any other change; at other times attended by +more serious alterations. So far as can be judged from exceptional +occurrences of this kind, it would appear that in this order the axis or +flower-stalk does not, in any material degree, enter into the +composition of the fruit. + +In the Rubiaceous genus _Bikkhia_, as mentioned by Duchartre, the ovary +is completely inferior, but when the fruit arrives at maturity four +small leaves are detached from its surface which had previously adhered +to it, and which it seems reasonable to consider as the sepals. + +In _Campanulaceae_ a similar separation of calyx from the ovary may be +occasionally met with. On the other hand, the occasional formation of a +leaf on the inferior ovary of those plants would indicate the axial +nature of the fruit. In _Campanumaea_ and _Cyclodon_ the calyx is +inferior, while the corolla is superior. In the last-named genus this +peculiarity "is carried to the highest degree, the sepals being, in _C. +parviflorum_, placed on the peduncle of the flower far removed from the +base of the corolla and ovary, whilst in _C. truncatum_ and in +_Campanumaea_ they adhere to the base of the tube of the corolla."[87] In +this order, then, as in _Saxifragaceae_, _Bruniaceae_, &c., no hasty +conclusion should be drawn as to the nature of the fruit. In _Brunia +microphylla_ the ovary is superior, enclosed within but not adnate to +the cup-like calyx, to which latter, however, the petals and stamens are +attached. + +In _Onagraceae_ (_Jussieua_), as also in _Cactaceae_ (_Opuntia_), buds +have been observed on the surface and edges of the inferior ovary. +Indeed, in the former genus, they have been produced artificially, but +as buds may be formed on foliar as well as on axial organs, the fact +cannot be made great use of in support either of the foliar or axial +nature of the inferior ovary. In _Epilobium_, I have met with four +perfect leaves at the summit of the ovary, in the place usually occupied +by the sepals. This would also favour the notion that the axis entered +into the constitution of the fruit in this genus. + +Mr. B. Clarke, in his 'New Arrangement of Phanerogamous Plants,' p. 4, +cites a case wherein the perianth was completely detached from the +surface of the ovary in _Cannabis sativa_. + +It must be borne in mind that some of the recorded instances of change +in the relative position of the calyx and pistil ought more properly to +be referred to a substitution of carpels for stamens, as in _Begonia_, +_Fuchsia_, &c. Among _Cucurbitaceae_, examples have been recorded, both +of the detachment of the calyx from the ovary,[88] and of the partial +conversion of some of the anthers of the male flower to carpels. + +The very singular mode of germination of _Sechium edule_ in which the +fruit, instead of rotting, becomes thickened into a kind of rhizome or +tuber, is a fact that should not be overlooked in investigating the true +nature of the fruit in this order. + +The following are the genera in which the change has been most +frequently observed: + + *Rosa! + *Pyrus! + *Crataegus! + *Daucus! + Pastinaca. + Torilis. + Apium. + OEnanthe! + Heracleum, + Athamanta. + Selinum. + Carum. + Imperatoria. + Rudbeckia! + *Campanula! + Lonicera! + Cucumis! + Cannabis. + +=Solution of the stamens from the petals.=--A separation of the stamens +from the petals in flowers, wherein those organs are usually adherent +one to the other does not often occur unattended by other changes. It +has been observed in _Cobaea scandens_ (Turpin), in _Antirrhinum majus_, +and in many double flowers. + +Partial detachment of the stamens from the styles occurs frequently in +semi-double flowers of _Orchis_.[89] + +FOOTNOTES: + +[85] 'Gard. Chron.,' 1865, p. 554; 1867, p. 599. + +[86] 'Bull. Soc. Bot. Fr.' 1854, p. 303. + +[87] Hook et Thoms, 'Praecurs. ad Flor. Ind.,' Journ. Linn. Soc., vol. +ii, 1858, p. 6. + +[88] Lindley, 'Veget. Kingd.,' p. 315. + +[89] Masters, 'Journal of Linnean Society,' 1866, vol. viii, p. 207. On +the subject of this chapter the reader should also consult +Moquin-Tandon, 1. c., p. 298. Engelmann, 'De Antholysi,' p. 37, tab. v. +C. Morren, 'Bull. Acad. Belg.,' xix, part 3, p. 318. Cramer, +'Bildungsabweichungen,' p. 64. Fleischer, 'Missbild. Cultur. Pflanzen.' +As to the nature of inferior ovaries, see also Payer, 'Bull. Soc. Bot. +Fr.' i, 1854, p. 283. Germain de Saint Pierre, _ibid._, p. 302. Caspary, +'Bull. Soc. Bot. Fr.,' t. vi, 1859, p. 235. Schleiden, 'Principles of +Botany,' English translation, p. 368. Duchartre, 'Elements de +Botanique,' p. 574. Le Maout et Decaisne, 'Traite general de Botanique,' +p. 57. Bentham, 'Journ. Linn. Soc.,' vol. x, p. 104 (Structure of +_Myrtaceae_), and other treatises on Organography. + + + + +PART III. + +ALTERATIONS OF POSITION. + + +Necessarily connected with changes in the arrangement of organs are +similar alterations in their position; so closely, indeed, that but for +convenience sake, it would be unnecessary to treat them separately. +There are, however, some anomalous developments affecting the relative +position of organs that could hardly be treated of under any of the +preceding paragraphs. There are, also, certain rare instances where an +organ is not so much displaced as misplaced; that is to say, it is +developed on or from a portion of the plant, which under usual +circumstances does not produce such an organ. In the former instance, +the altered position is due to or coexistent with other changes, but in +the latter case the new growth may spring from organs otherwise in +nowise different from ordinary. The word Displacement is here used to +signify the unusual position of an organ; while Heterotaxy may serve to +include those cases where a new growth makes its appearance in an +unwonted situation, as, for instance, a leaf-bud on a root, &c. +Prolification is also included under this heading, the unusual position +of the buds in these cases being of graver import than the mere increase +in number. Alterations in the position of the sexual organs are spoken +of under the head of Heterogamy. + + + + +CHAPTER I. + +DISPLACEMENT. + + +Real or apparent displacement of organs from their usual position is an +almost necessary consequence of, or is, at least, coexistent with a +large number of teratological phenomena. It is obvious that abnormal +unions or disunions, suppressions, hypertrophies, &c., are very liable +to bring about or to be accompanied with changes in the position, either +of the parts directly affected or of adjoining organs. + +In this place, then, it is merely necessary to allude to some of the +more important displacements, and to refer for further details to the +sections relating to those irregularities of growth on which the +displacement depends. + +=Displacement of bulbs.=--I owe to the kindness of Mr. James Salter a +tulip bulb which had been dug up after flowering, and from the base of +which were suspended several small bulbs; and I have since seen another +specimen showing the same unusual arrangement. The explanation of these +formations seems to be that they correspond to the bulbils ordinarily +found in the axils of the scales of the parent organ, and which, in some +way or another, have been displaced and thrust into the ground. +Professor de Vriese figures something of the same kind in _Ixia +carminosa_.[90] + +Of somewhat different nature to those above described was an anomaly +described by M. Gay at a meeting of the Botanical Society of France, +April 8th, 1859. The plant affected was _Leucoium aestivum_, and the +changes observed were apparently attributable to a simple separation of +two leaves that are usually contiguous. "Suppose," says M. Gay in +describing this malformation, "the first leaf of the terminal bud +separated by a long internode from the other leaves, which remain +closely packed; and further, suppose an evident thickening of the upper +portion of the lengthened internode, and there will be not only a single +bulb, bearing with the leaves of the present year all the remnants of +the leaves of the two preceding years, but two bulbs placed one above +another, on the same axis, separated by the length of the internode." + +[Illustration: FIG. 39.--Unusual position of bulbs of tulip; the +parent-bulb cut open.] + +The formation of bulbs in the axils of the leaves, as happens +occasionally in tulips, is further alluded to under the head of +hypertrophy. + +=Displacements affecting the inflorescence.=--These are, for the most +part, dependent on hypertrophy, elongation, atrophy, spiral torsion, +&c., but there are a few instances of a different nature, which may here +be alluded to as not being coincident with any of the phenomena just +mentioned. Sometimes these deviations from the ordinary position have +the more interest as affecting characters used to distinguish genera; +thus one of the distinctions between rye-grass (_Lolium_) and wheat +(_Triticum_) resides in the relative position of the spikelets and the +main stem; in _Triticum_ the spikelets are placed with their backs +against the rachis, in _Lolium_ with one edge against it; but in a +specimen of rye-grass that has come under my own observation, the +arrangement was that of _Triticum_. + +M. Kirschleger relates having found a specimen of _Leucanthemum +pratense_, in which the ligulate female flowers were growing singly in +the axils of the upper leaves of the stem.[91] The ordinary capitulum +would here seem to have been replaced by a spike or a raceme. A less +degree of this change wherein a few flowers may be found, as it were, +detached from the ordinary capitulum may often be observed in +_Compositae_, _Dipsacaceae_, &c. I have also met with specimens of _Lamium +album_ in which some of the fascicles or clusters of flowers in place of +being placed at the same level on opposite sides of the stem were placed +alternately one above another. + +Caspary[92] mentions a flower of _Aldrovanda vesiculosa_, which was +elevated on a stalk that was adherent to the stem for a certain +distance, and then separated from it. This flower, with the leaf to +which it was axillary, evidently belonged to the whorl beneath, where +there was a corresponding deficiency. Another flower of the same plant +bore on its pedicel a small leaf, which was doubtless the bract raised +above its ordinary position. + +M. Fournier mentions an instance in _Pelargonium grandiflorum_, where, +owing to the lengthening of the axis, the pedicels, instead of being +umbellate, had become racemose; and I owe to the kindness of Dr. Sankey +a somewhat similar specimen, but in a less perfect condition. Here there +was but a single flower, and that rudimentary, placed at the extremity +of the axis. There were several bracts beneath this flower disposed +spirally in the 1/3 arrangement, all being empty, excepting the terminal +one. In like manner, a head of flowers becomes sometimes converted into +an umbel. + +=Displacement of leaves.=--A cohesion of parts will sometimes give rise +to an apparent displacement, but the true nature of the malformation +can, in general, be readily made out. + +Steinheil[93] found a specimen of _Salvia Verbenaca_, the leaves of +which presented very curious examples of displacement arising from +cohesion. Two of these leaves placed at the base of a branch were +completely fused in their lower thirds, and divided into two distinct +lobes at the upper part; each of these lobes seemed to be as large as +the limb of an ordinary leaf. Above these was another very broad one, +apparently entire, but evidently produced by a complete cohesion of two. +This completely fused leaf alternated in position with the imperfectly +fused one below it; the alternation is explained by supposing that the +opposite leaves of each pair were directed one towards the other, and +became fused, and that thus resulted the displacement. The dislocation +of the organs took place in one direction for one pair of leaves, and in +another direction for the other pair, hence the alternation. Thus, +leaves normally opposite and decussate may, by fusion, become alternate. +A similar instance occurred to the writer in _Lysimachia vulgaris_, +wherein the changes arising from fusion and suppression of parts, &c., +were very considerable; as far as the leaves were concerned they +presented the following arrangement in succession from below +upwards:--first verticillate, then opposite, then spirally alternate, +lastly opposite.[94] The term "diremption" has sometimes been applied to +cases where leaves are thus apparently dragged out of position. + +In _Tradescantia virginica_ I have met with opposite connate leaves; +the altered position, however, being due to the union of two stems. + +[Illustration: FIG. 40.--Large-coloured leaf occupying the position of +the inflorescence in _Gesnera_, after Morren.] + +[Illustration: FIG. 41.--Ordinary arrangement of leaves in fascicles of +three in _Pinus pinea_ and unusual arrangement of leaves of same plant +in spires.] + +Twisting of the stem is a frequent cause of the displacement of leaves +(see spiral torsion), as also hypertrophy, whether that excess of +development take place laterally or lengthwise (see elongation). Atrophy +or suppression will also frequently bring about an alteration in the +position of leaves; sometimes in such a manner that the place of the +suppressed organ is occupied by another one. One of the most curious +instances of displacement of leaves arising from suppression is that +mentioned by Morren,[95] where, in _Gesnera Geroltiana_, a large leaf +apparently occupied the extremity of the axis, a position which, under +ordinary circumstances, no leaf could assume. The explanation given by +the Belgian professor is, that the axis in this case, instead of +throwing off a pair of leaves, one on each side, had from some cause or +another produced only one; this one not only being much larger than +ordinary, but brightly coloured, thus assuming some of the +characteristics as well as the position of the inflorescence. + +Alterations in the usual arrangement of leaves, however, are not always +dependent on or coexistent with other teratological changes, but may +simply depend on a natural elongation of the internodes, or on fission +or multiplication; for instance, in some conifers, such as the Larch, +(_Abies Larix_) or _Pinus pinea_, there may be found at different stages +in the growth of the branches leaves in crowded fascicles or tufts; +while, when growth is more rapid, the leaves may be disposed in a spiral +or alternate manner. + +In the yew (_Taxus_) the leaves at the ends of the shoots not +unfrequently lose their usual distichous arrangement and become arranged +in a close spiral manner, the elongation of the shoot being arrested. +This appears to be the result of the injury effected by some insect. + +[Illustration: FIG. 42.--Altered arrangement of leaves of yew, _Taxus +baccata_.] + +So, too, the alteration from verticillate to spiral, or _vice versa_, +may take place without any other notable change.[96] This may frequently +be seen in Rhododendrons. + +=Displacement of the parts of the flower.=--This subject is partly +touched on in the chapters on solution, adhesion, and in those on +hypertrophy, elongation, prolification, &c., so that in this place it is +only requisite to offer a few general remarks, and to refer to other +sections for further details. Morren, in referring to displacement of +the floral organs, mentions an instance in a _Fuchsia_, wherein the four +petals in place of being alternate with the sepals were placed in front +of them, owing to the adhesion that had taken place between the petals +and the stamens. He speaks of this transposition as metaphery.[97] The +same author also gives an account of the displacement of several of the +organs of the flower in _Cypripedium insigne_, the displacement being +consequent, apparently, on a spiral torsion proceeding from right to +left, and involving the complete or partial suppression of several of +the organs of the flower. The dislocation of organs in a spiral +direction led Morren to apply the term "speiranthie" to similar +deviations from the usual construction. Changes of this kind among +_Orchidaceae_ are by no means uncommon; the following may be cited by way +of illustration. In a specimen of _Oncidium cucullatum_ furnished me by +Mr. Anderson, well known for his success as a cultivator of these +plants, there was, associated with a cohesion of one sepal with another, +and probably dependent on the same cause, a displacement of the sepals +and petals--so that all were dragged out of place. This dislocation may +be better appreciated by the accompanying formula than even by the +woodcut. Let the usual arrangement be thus represented: + + S + + P ST P + + L + + S S + +S standing for sepal, P for petal, L for lip, ST for stamen; then the +dislocated form may be represented thus: + + S + + P P + + T + S + + S_S + L + +[Illustration: FIG. 43.--Flower of _Oncidium cucullatum_, showing union +of two lower sepals, displacement of column and lip, &c.] + +In a specimen of _Cypripedium_ also furnished by Mr. Anderson the +appearance was as represented in the accompanying figure and diagrams, +figs. 44, 45. Referring to the plan of the natural arrangement at fig. +46, it will be seen that an explanation of the peculiar appearance of +the flower may be arrived at by supposing a disunion and lateral +displacement of the upper segment of the outer perianth together with +the complete absence of the lower one. In the second or inner whorl of +the perianth the lip is merely a little oblique on one side, but the +lateral petals are distorted, displaced, and adherent one to the other +and to the column, while the posterior shield-like rudimentary anther is +completely wanting. + +[Illustration: FIG. 44.--Malformed flower of _Cypripedium_.] + +[Illustration: FIG. 45.--Diagram of malformed _Cypripedium_. _o_, outer +segments; _i_, inner segments of perianth; _e_, lip; _s_, stigma; _a_, +anther.] + +[Illustration: FIG. 46--- Diagram showing ordinary arrangement in +_Cypripedium_. _o_, outer, _i_, inner segments of perianth; _e_, lip, +_a_, anther, _a'_, abortive stamen; _s_, stigma.] + +[Illustration: FIG. 47.--Plan of flower of _Lycaste Skinneri_ showing +displacement of organs.] + +In a specimen of _Lycaste Skinneri_ similar changes were observed, as +shown in the plan, fig. 47. Here the posterior sepal was deficient, the +two lateral ones were present, one of them with a long tubular spur, _o +o_; of the two lateral petals, _i i_, one was twisted out of place, so +as partially to occupy the place of the deficient sepal; the lip was +represented by two three-lobed segments, _l_, one above and within the +other. The column and ovary of this flower were in their normal +condition. + +Cohesion of two or more segments of the perianth is frequently +associated with displacements of this nature: thus, in a flower of +_Dendrobium nobile_, a diagram of which is given at fig. 48, the +uppermost sepal was coherent with one of the lateral ones, and at the +same time diminished in size, and, as it were, dragged out of position. +All the other organs of the flower are also more or less displaced, +forming a minor degree of the change already alluded to, and which +Morren termed speiranthy. The changes will be better appreciated by +comparing them with fig. 49, a diagram showing the natural arrangement +of parts in this species. + +[Illustration: FIG. 48.--Plan of malformed flower of _Dendrobium +nobile_.] + +[Illustration: FIG. 49.--Plan of natural arrangement in _Dendrobium +nobile_. The x x represent processes of the column, perhaps rudiments of +stamens.] + +Sometimes the displacement seems consequent on hypertrophy of one of the +parts of the flower, the disproportionate size of one organ pushing the +others out of place. This was the case in a violet, fig. 50, in which +one of the sepals _s_ was greatly thickened, and the petals and stamens +were displaced in consequence. + +[Illustration: FIG. 50.--Plan of flower of violet showing displacement +of petals, &c. At _b_ was a rudiment of a stamen.] + +It is curious to observe in many of these cases that the transposed +organ not only occupies the place of a suppressed or abortive organ, +but frequently assumes its colour, and, to some extent, its function. +This has been alluded to in the case of the leaf of _Gesnera_ (see p. +88) and in Orchids this replacement seems to be very common; thus, in +addition to the cases before mentioned, in a flower of an Odontoglossum, +for which I am indebted to Professor Oliver, the two lateral sepals were +united together and occupied the position of the labellum, which was +absent. A similar occurrence happens occasionally in _Lycaste Skinneri_, +thus recalling the structure of _Masdevallia_, where the labellum is +normally very small. The arrangement in Lycaste may thus be symbolised: + + S + P st P + + + S S + ___ + +[Transcriber's note: The underscores represent a horizontal curly brace +in the original.] + +the + indicating the position of the absent labellum. + +Cases of this kind are the more interesting from their relation to the +fertilization of these flowers by insects; it seems as though, when the +labellum, which performs so important an office in attracting and +guiding insects, is deficient, its place is supplied by other means. + +Displacement of the parts of the flower from elongation of the +receptacle is a not infrequent teratological occurrence, resulting +sometimes in the conversion of the verticillate into the spiral +arrangement. Instances of this are cited under Elongation, +Prolification, &c. In this place it is merely necessary to refer to a +curious circumstance that is met with in some double flowers, owing to +this separation of some parts of the flower and the cohesion or adhesion +of others. Thus, in some double flowers of _Primula sinensis_ and in the +Pea (_Pisum sativum_), I have seen a gradual passage of sepals to +petals, so that the calyx and corolla formed one continuous sheet, +winding spirally around the central axis of the flower, after the +fashion of a spiral tube.[98] + +=Displacement of the carpels= arises from one or other of the causes +above alluded to, and when suppression takes place in this whorl it +generally happens that the place of the suppressed organ is occupied by +one of the remaining ones, which thus becomes partially dislocated. + +=Displacement of the placentas and ovules= is a necessary result of many +of the changes to which the carpels are subject. The disjunction or +dialysis of the carpels, for instance, frequently renders axile +placentation marginal. Moreover, it frequently happens, when the carpels +become foliaceous and their margins are disconnected, that the ovules, +in place of being placed on the suture, or rather on the margins of the +altered carpel, are placed on the surface of the expanded carpel. Thus, +in some double flowers of _Ranunculus Ficaria_ that came under the +writer's notice the carpels were open, _i.e._ disunited at the margins, +and each bore two imperfect ovules upon its inner surface a little way +above the base, and midway between the edges of the carpel and the +midrib, the ovules being partly enclosed within a little depression or +pouch, similar to the pit on the petals. On closer examination the +ovules were found to spring from the two lateral divisions of the +midrib, the vascular cords of which were prolonged under the form of +barred or spiral fusiform tubes into the outer coating of the ovule. In +this instance, then, the ovules did not originate from the margins of +the leaf, nor from a prolonged axis, but they seemed to spring, in the +guise of little buds, from the inner surface of the carpellary leaf.[99] + +The occurrence, also, of different forms of placentation in different +flowers on the same plant is no unusual thing in malformed flowers; +thus, in double flowers of _Saponaria officinalis_ I have met with +sutural, parietal, and free central placentation in the same plant.[100] + +Professor Babington describes in the 'Gardeners' Chronicle,' 1844, p. +557, a curious flower of _Cerastium_, in which, in addition to other +changes, the five carpellary leaves "were partially turned in without +touching the placenta, which bears a cluster of ovules, and is perfectly +clear of all connection with those partitions" (fig. 51). See also +Lindley, 'Veg. Kingdom,' p. 497. + +[Illustration: FIG. 51.--1. Monstrous flower of a _Cerastium_; sepals +and petals leafy. 2. Stamens and pistils separate. 3. Ovary cut open to +show the imperfect dissepiments and the attachment of the ovules. 4. A +deformed ovule.] + +M. Baillon[101] records flowers of _Bunias_, some with ovules on the +margins of the carpels, others with a central branch bearing the ovules; +hence he concludes very justly that no fair inference can be drawn from +these facts as to the normal placentation of Cruciferae. + +The same excellent observer has recorded the occurrence of free central +placentation in malformed flowers of _Trifolium repens_.[102] + +In malformed flowers of _Digitalis_ the change from axile to parietal +placentation may often be seen. Mr. Berkeley describes an instance of +this nature where the placentas were strictly parietal, and therefore +receded from the distinctive characters of the order, and approximated +to those of _Gesneraceae_. + +The same author alludes to certain changes in the same flower where two +open carpels "were soldered together laterally, as was clear by the +rudiments of two styles, the placenta being produced only at the two +united edges, the outer margins remaining in the normal condition. This +may possibly tend to the explanation of some cases of anomalous +placentation, for the only indication of the true nature of the +placentation is afforded by the two rudimentary styles, in the absence +of which the spongy receptacle of the seeds must have been supposed to +spring from the medial nerve." + +In other cases the placentas were parietal above, but axile at the base +of the capsule, a striking instance of the facility with which axile +placentation becomes parietal, the change being here effected by the +prolongation of the axis, and the formation on it of a second whorl of +carpellary leaves. + +In double flowers of _Primulaceae_ similar alterations in the +placentation may often be observed. I have seen in _Primula sinensis_ +sutural, parietal, axile, and free central placentation all on the same +plant; nay, even in the same capsule the ovules may be attached in +various ways, and transitions from one form of placentation to another +are not infrequent. The late Professor E. Forbes describes[103] an +instance of true foliar and true axile placentation in the same flower +in _Vinca minor_. + +These and many similar changes, which it is not necessary further to +allude to, are not so much to be wondered at when it is borne in mind +how slight an alteration suffices to produce a change in the mode of +placentation, and how frequent is the production of adventitious buds or +of foliar outgrowths, as may be seen in the sections relating to those +subjects and to Substitutions. + +It will be remembered, also, how, in certain natural orders, under +ordinary circumstances, considerable diversity in placentation exists, +according as the margins of the carpels are merely valvate or are +infolded so as to reach the centre. Often this diversity is due merely +to the changes that take place during growth; thus, the placentation of +_Caryophylleae_, _Cucurbitaceae_, _Papaveraceae_, and many other orders, +varies according to the age of the carpel, and if any stasis or arrest +of development occurs the placentation becomes altered accordingly. + +It is not necessary, in this place, to enter into the question whether +the placenta is, in all cases whatsoever, a dependence of the axis, as +Payer, Schleiden, and others, have maintained, or whether it be foliar +in some cases, axial in others. This question must be decided by the +organogenists; teratologically, however, there can be no doubt that +ovules may be formed from both foliar and axial organs, and, moreover, +that, owing to the variability above referred to, both in what are +called natural and in what are deemed abnormal conditions, it can rarely +happen that any safe inferences as to the normal or typical placentation +of any family of plants can be drawn from exceptional or monstrous +formations. + +On the subject of placentation the following authors may be consulted: + + R. Brown, 'Ann. Nat. Hist.,' 1843, vol. xi, 35. Brongniart, + 'Ann. Sc. Nat.,' 1834, ser. 2. i, p. 308. Alph. De Candolle, + 'Neue Denkschrift der Allg. Schweizer Gesellsch.,' Band v. + 1841, p. 9. Duchartre, 'Ann. Sc. Nat.,' 3rd ser., 1844, vol. + ii, p. 290. Ibid., 'Elem. Bot.,' p. 574; 'Rev. Bot.,' 1846-7, + p. 213. Babington, 'Gard. Chron.,' 1844, p. 557. Lindley, + 'Elements,' p. 89; 'Veg. King.,' pp. 313, 497, &c. Berkeley, + 'Gard. Chron.,' 1850, p. 612. Unger, 'Nov. Act. Acad. Nat. + Cur.,' 1850; and in Henfrey's, 'Bot. Gazette,' 1851, p. 70. + Schleiden, 'Principles,' English edit., p. 385. Payer, 'Elem. + Bot.,' pp. 196, 211, 224. Baillon, 'Adansonia.' iii, p. 310. + tab. iv. Cramer, 'Bildungsabweichungen,' p. 20, &c. Clos, 'Ann. + Sc. Nat.,' 5th ser., iii, 313, as well as any of the general + treatises on botany. Reference may also be made to the chapters + on Prolification and Substitutions (in the case of the carpels + and ovules), and to the authorities therein cited. + +FOOTNOTES: + +[90] 'Tijdschr. voor. nat. Gesch.,' viii, 1841. tab. ii, p. 178. + +[91] Communication to the Internat. Bot. Congress, Paris, 1867. + +[92] 'Bot. Zeit.,' 1859, p. 117, tab. v. + +[93] 'Ann. Sc. Nat.,' ser. 2, vol. iv, 1835, p. 143. tab. v. + +[94] See Kirschleger, 'Flora.' 1844. p. 566 (_Scabiosa_). + +[95] 'Bull. Acad. Belg.,' t. xvii. part ii, p. 387. + +[96] 'Clos. Mem. Acad. Toulouse,' 5th ser., t. vi. pp. 51, 70. + +[97] 'Bull. Acad. Roy. Belg.,' xviii. part ii, p. 505, and vol. xvii, +part i, p. 196, and vol. xix. part i. p. 260. + +[98] See also Schlechtendal, 'Bot. Zeit.,' iv, p. 804. _Primula veris, +partibus perigonii spirae in modum confluentibus._ + +[99] Seemann's 'Journal of Botany,' vol. v, 1867, p. 158. + +[100] 'Journ. Linn. Soc.,' i, 1857, p. 161. _c. xylog._ + +[101] 'Adansonia,' ii, 306. + +[102] 'Adansonia,' iv, p. 70, t. i. + +[103] Henfrey's 'Bot. Gazette,' i, 265. + + + + +CHAPTER II. + +PROLIFICATION. + + +Moquin-Tandon and other writers have classed the production of buds in +unwonted situations under the head of multiplication, but, as the +altered arrangement is of graver import than the mere increase in +number, it seems preferable to place these cases under this heading +rather than under that of alterations of number. + +The adventitious bud may be a leaf-bud or a flower-bud; it may occupy +the centre of a flower, thus terminating the axis, or it may be axillary +to some or other of its component parts, or, again, it may be +extra-floral. In this last case the prolification is of the +inflorescence, and is hardly distinguishable from multiplication or +subdivision of the common flower-stalk. In accordance with these +differences we have median, axillary, and extra-floral prolification, +each admitting of subdivision into a leafy or a floral variety, +according to the nature of the adventitious bud. Under the head of each +variety certain special peculiarities are noticed, but it may here be +advisable to add a few general remarks on the subject. + +Axillary prolification is a much less frequent malformation than the +median form. If only the number of orders and genera be reckoned, the +truth of this statement will be scarcely recognised; but if individual +cases could be estimated, the difference in frequency between the two +would be very much more obvious. This may, perhaps, be explained by the +fact that the branch has a greater tendency to grow in length than it +has to develop buds from the axils of the leaves. The flower is admitted +to be homologous with the branch, and it is also known that, up to a +certain time, the branch-bud or leaf-bud and the flower-bud do not +essentially differ.[104] At a later stage the difference between the two +is manifested, not only in the altered form of the lateral organs in the +flower-bud, but in the tendency to an arrest of growth, thus limiting +the length of the central axial portion. Now, in prolified flowers the +functions and, to a considerable extent, the appearance of a leaf-bud or +of a branch are assumed, and with them the tendency to grow in length is +developed. Median prolification, therefore, in this sense, is a further +step in retrograde metamorphosis than is the axillary form. To grow in +length, and to produce axillary buds, are alike attributes of the +branch; but the former is much more frequently called into play than the +latter; for the same reason, median prolification is more common than +the axillary form. This is borne out by the frequency with which +apostasis, or the separation of the floral whorls one from another, to a +greater degree than usual, is met with in prolified flowers. + +In both forms the adventitious growth is much more frequently a +flower-bud or an inflorescence than a leaf-bud or a branch. This may be +due to the position of the flowers on a portion of the stem of the plant +especially devoted to the formation of flower-buds, to the more or less +complete exclusion of leaf-buds, _i.e._ on the inflorescence. This +conjecture is borne out by the comparative rarity with which +prolification has been observed in flowers that are solitary in the +axils of the ordinary leaves of the plant. If the lists of genera +appended hereto be perused, it will be seen that nearly all the cases +occur in genera where the inflorescence is distinctly separated from the +other branches of the stem. In direct proportion, then, to the degree in +which one region of the axis or certain branches of a plant are devoted +to the formation of flower-buds to the exclusion of leaf-buds, is the +frequency with which those flowers become affected with floral +prolification. + +Flowers produced upon indefinite inflorescences are liable to be +affected with either form of prolification more frequently than those +borne upon definite inflorescences. Prolification in both varieties is +also more frequently met with in branched inflorescences than in those +in which the flowers are sessile; but the degree of branching seems less +material, inasmuch as this malformation is more commonly recorded as +occurring in racemes than in the more branched panicles, &c. From the +similar arrest of growth in length, in the case of the flower, to that +which occurs in the stem in the case of definite inflorescence, it might +have been expected that axillary prolification would be more frequent in +plants having a cymose arrangement of their flowers than in those whose +inflorescence is indefinite; such, however, is not the case. The reason +for this may be sought for in the lengthening of the floral axis, so +common in prolified flowers--a condition the reverse of that which +happens in the case of definite inflorescence. + +Median prolification occurs frequently in double flowers; the axillary +variety, on the other hand, is most common in flowers whose lateral +organs have assumed more or less of the condition of leaves. The other +coincident changes are alluded to elsewhere or do not present useful +points of comparison, and may therefore be passed over. + +=Prolification of the inflorescence.=--This consists in the formation of +leaf-buds or of an undue number of flower-buds on the inflorescence. It +must be distinguished from virescence, or the mere green colour of the +floral organs, and from chloranthy, in which all or the greater portion +of the parts of the flower are replaced by leaves. Prolification is, in +fact, a formation of supernumerary buds, leafy or floral, as the case +may be, these buds being sessile or stalked, the ordinary buds being not +necessarily changed. Prolification of the inflorescence, like the other +varieties, admits of subdivision, not only according to the foliar or +floral nature of the bud, but according to its position, terminal or +median and lateral. + +Terminal prolification of the inflorescence, whether leafy or floral, is +hardly to be looked upon in the light of a malformation[105] seeing that +a similar condition is so commonly met with normally, as in _Epacris_, +_Metrosideros_, _Bromelia_, _Eucomis_, &c., wherein the leafy axis +projects beyond the inflorescence proper; or as in _Primula imperialis_, +in which plant, as also in luxuriant forms of _P. sinensis_, tier after +tier of flowers are placed in succession above the primary umbel. +Nevertheless, when we meet with such conditions in plants which, under +ordinary circumstances, do not manifest them, we must consider them as +coming under the domain of teratology. + +=Median foliar prolification of the inflorescence= is frequently met +with in _Coniferae_, and has of late attracted unwonted attention from +the researches of Caspary, Baillon, and others, on the morphology of +these plants. The scales and bracts of the cone in these abnormal +specimens frequently afford transitional forms of the greatest value in +enabling morphologists to comprehend the real nature of the floral +structure. It would be irrelevant here to enter into this subject; +suffice it merely to say that an examination of very numerous specimens +of this kind, in the common larch and in _Cryptomeria Japonica_, has +enabled me to verify nearly the whole of Caspary's observations. A +similar prolongation of the axis occurred in some of the male catkins of +_Castanea vesca_, each of which had a tuft of small leaves at their +extremity. In the common marigold and in _Lotus corniculatus_ I have +also seen instances of this kind. Kirschleger[106] describes a tuft of +leaves as occurring on the apex of the flowering spike after the +maturation of the fruit in _Plantago_, and a similar growth frequently +takes place in the common wallflower, in _Antirrhinum majus_, &c. In +cases where a renewal of growth in the axis of inflorescence has taken +place after the ripening of the fruit, the French botanists use the term +recrudescence, but the growth in question by no means always occurs +after the ripening of the fruit, but frequently before. Professor Braun +cites the case of a specimen of _Plantago lanceolata_, in which the +spike was surmounted by a tuft of leaves and roots, as well as a still +more singular instance in _Eryngium viviparum_, in which not only did +particular branches terminate in rosettes of leaves provided with roots, +but similar growths proceeded from the heads of flowers themselves. +Baron de Melicoq[107] gives a case in _Primula variabilis_, in which at +the top of the flower-stalk, in the centre of six flowers, was placed a +complete plant in miniature, having three leaves, from the axil of one +of which proceeded a rudimentary flower. Mr. W. B. Jeffries also +forwarded me a polyanthus (fig. 52) in which the peduncle was surmounted +by a small plant, forming a crown above the ordinary flower-stalk, just +as the crown of the pineapple surmounts that fruit. A similar instance +was exhibited at the Scientific Committee of the Horticultural Society +on July 11th, 1868, by Mr. Wilson Saunders; the species in this case was +_P. cortusoides_. To Mr. R. Dean I am indebted for a similar proliferous +cyclamen, which seems similar to one mentioned by Schlechtendal.[108] +This author alludes to an analogous circumstance in the inflorescence of +_Cytisus nigricans_, where, however, the change was not so great as in +the preceding cases. The instances just cited all occur in plants having +an indefinite form of inflorescence; but the production of a tuft of +leaves or of a leafy shoot above or beyond the inflorescence is not +confined to plants with this habit of growth, for Jacquin figures and +describes an instance of this nature in the cymose flower-stems of a +Sempervivum. "_Hi racemi_," says he, "_ultra flores producuntur in +ramos, foliosos duo bifidos qui tandem trium unciarum longitudinem +adepti fuerunt_."[109] + +[Illustration: FIG. 52.--Inflorescence of _Polyanthus_, bearing a tuft +of leaves at the top of the scape intermixed with the flowers.] + +=Median floral prolification of the inflorescence=, wherein a new +inflorescence projects beyond the primary one, is not uncommon in plants +having their flowers arranged in close heads or umbels, as in the +common wild celery and other _Umbelliferae_.[110] I have also met with +it in _Trifolium repens_, in the umbellate variety of the common +primrose, and in the scarlet geranium. Engelmann cites it in _Triticum +repens_, Roeper in _Euphorbia palustris_.[111] + +=Lateral foliar prolification of the inflorescence= is of more common +occurrence than the preceding. I have met with it, amongst other plants, +frequently in _Brassica oleracea_, _Pelargonium zonale_, SCABIOSA, +BELLIS, and many other composites, also in _Leguminosae_, e.g. _Lupinus_, +_Trifolium_, _Coronilla_, &c. Prof. Oliver forwarded me a specimen of +_Euphorbia geniculata_ in which, in addition to other changes, there was +a series of stalked buds bearing tufts of green scales, but without any +trace of stamens or pistil; these adventitious buds occurred within the +ordinary involucre of the plant, between it and the stamens. The pistil +was unaffected in some cases, while in some others it was entirely +wanting, the gynophore being surmounted by a cup-like involucre, divided +into three acutely pointed lobes, each with a midrib; these encircled a +series of stalked involucels, as before, and among which were scattered +a few stamens, some perfect, others partially frondescent. + +In a specimen of _Scrophularia nodosa_ examined by me one of the lateral +buds on each of the cymes was represented, not by a flower, but by a +tuft of leaves, the other buds being unchanged. As the inflorescence was +much contracted in size, the appearance of the whole plant was greatly +changed. + +Many of the instances of so-called viviparous plants, _e.g._, _Polygonum +viviparum_, may be cited under this head.[112] Many species of _Allium_, +_Lilium_, _Saxifraga_, _Begonia_, _Achimenes_, normally produce +leaf-buds or bulbs in the inflorescence; so, too, leafy shoots are +sometimes found in _Alisma natans_, _Juncus uliginosus_, _Chlorophytum +Sternbergianum_, &c. As an accidental occurrence, a similar thing has +been noticed in _Lychnis coronaria_, _Phaius grandifolius_, _Oncidium +cebolleta_, _Epidendrum elongatum_,[113] &c. &c. + +Here, too, may be mentioned those cases wherein a leaf-bud is found upon +the surface of the so-called inferior ovary; generally a leaf only is +found, but a leaf-bud may also originate in this situation, and in +either case the inference is that the ovary is, in part at least, made +of the dilated and hollowed axis. Leaves may occasionally be found in +this way on the so-called calyx-tube or on the inferior ovaries of +roses, pears, apples, _Pereskia_, _Crataegus tanacetifolia_, &c. + +The fruits of _Opuntia Salmania_ and of _O. fragilis_ ('Bull. Soc. Bot. +France,' vol. i, p. 306; vol. v, p. 115) have been observed to form +small fruit-like branches around their summits. This circumstance is +more fully treated of in the succeeding chapter relating to Heterotaxy. + +=Lateral floral prolification of the inflorescence.=--This, which is +termed by Engelmann Ecblastesis foliorum sub floralium,[114] is much the +most common of all these deviations, and it is met with in every degree, +from the presence of a single supernumerary flower in the axil of a +bract to the existence of a small cluster or panicle of such flowers. + +[Illustration: FIG. 53.--Lateral prolification in inflorescence of +_Pelargonium_.] + +It is common in the _Anemone coronaria_ and _hortensis_, also in the +common scarlet _Pelargonium_ (fig. 53). It has been frequently recorded +in _Poterium sanguisorba_, and in _Sanguisorba officinalis_, and is +especially common in _Umbelliferae_, _Dipsacaceae_, and _Compositae_; a +familiar illustration in the latter order is afforded by the +hen-and-chicken daisy. In some species of Compositae, indeed, it is a +normal and constant occurrence, while in other cases, such as _Filago +germanica_, usually described as proliferous, there is not, strictly +speaking, any prolification, for the branching of the stalk takes place +below the inflorescence, and the branches originate from the axils of +ordinary leaves, not from the floral leaves or bracts. _Convolvulus +Sepium_ is very commonly subject to the production of flower-buds from +the axils of the floral leaves. The several species of Plantain +(_Plantago_) seem very liable to this and similar changes. +Schlechtendal[115] gives a summary of the various kinds of malformation +affecting the inflorescence in _Plantago_, and divides them into five +groups, as follows:--1st, bracteate, wherein the inferior bracts are +quite leaf-like, as is frequently seen in _Plantago major_. 2nd, +roseate; bracts leafy in tufts or rosettes, without flowers, as in the +so-called rose plantain, common in old-fashioned gardens in this +country. 3rd, polystachyate; spike-branched, bearing other spikes in the +axils of the bracts, as in _P. lanceolata_, _P. maritima_, &c. 4th, +proliferous, where the flower-stalk bears a rosette, a spike, or a head +with other rosettes. 5th, paniculate, in which the inflorescence has +become a much-branched pyramidal panicle, covered with little bracts, +and with very rudimentary flowers.[116] The first two groups belong +rather to frondescence of the bracts; but with regard to the whole of +them it will easily be surmised that intermediate forms occur, linking +one group to the other, and defying exact allocation in either. Thus, in +the borders of richly cultivated fields in the neighbourhood of London I +have frequently gathered specimens of _Plantago major_ with a branched +spike provided with large leafy bracts, the branches of the spike being +but little less in diameter than the ordinary single spike. These +specimens would therefore seem to be intermediate between +Schlechtendal's bracteate and polystachyate divisions. Wigand[117] also +describes an anomalous specimen of _Plantago major_ similar to those +just mentioned, but having small lateral spikes in place of large ones. +The instance quoted from Professor Braun would fall under the roseate +section, as would also that of Kirschleger, though we are expressly told +that the tuft of leaves in this last case was not developed until after +the ripening of the seed-vessel. One of the characters of the roseate +group, according to Schlechtendal, is the absence of flowers, but most +persons who have had the opportunity of watching the growth of the rose +plantain must have observed the occasional production of flowers, +sometimes stalked, in the axils of the leafy bracts, and at the same +time have noticed that the internodes become elongated, so that an +approach is made to the ordinary spike-like form of the inflorescence. +The proliferous group would include such specimens as that of _P. +lanceolata_ mentioned by Dr. Johnston,[118] wherein were several +spikes, some sessile, others stalked and pendent, the whole intermixed +with leaves and disposed in a rose-like manner. I have myself gathered +specimens of this nature, occurring in the same plant, at Shanklin, Isle +of Wight (fig. 56). + +[Illustration: FIG. 54.--_Plantago major_, with panicled inflorescence.] + +[Illustration: FIG. 55.--Inflorescence of _Plantago major_, with bracts +partly replaced by leaves and spike branched.] + +[Illustration: FIG. 56.--Inflorescence of _Plantago lanceolata_, bearing +a tuft of leaves and flowers at the end of the flower-scape.] + +It is rather singular that each species of _Plantago_ seems to have its +own perverse mode of growth; for instance, the bracteate, polystachyate +and paniculate forms are almost exclusively confined to _P. major_, the +roseate form to _P. media_, the proliferous form to _P. lanceolata_. + +The instances wherein flower-buds originate from the surface of an +inferior ovary, as in those cases where the top of the stem is dilated +so as to form part of the fruit, would be properly classed under the +head of prolification of the inflorescence. As, however, there is still +some difference of opinion as to the correct morphological +interpretation to be put on some of these cases, it has been thought +better to include them under the head of heterotaxy than of +prolification. + +[Illustration: FIG. 57.--Branched inflorescence of _Reseda luteola_.] + +Some of the cases of prolification of the inflorescence resulting in a +branching of an ordinarily simple inflorescence, as in _Reseda luteola_ +(fig. 57), might equally well be placed with fission or multiplication +of the axile organs. Branched spikes of this character are not so common +among Orchids as might be expected. Professor Reichenbach enumerates a +few instances in the Report of the International Botanical Congress of +London, 1866, p. 121, and the same author gives an illustration in his +'Orchidographia Europoea,' tab. 150. + +In Grasses, as indeed in other plants with a spicate inflorescence, this +change occurs not unfrequently. The common Ray Grass (_Lolium_) is +especially subject to the change in question, and among cultivated +cereals, maize and wheat occasionally show this tendency to subdivision. +One variety of the latter grain is cultivated in hot countries under the +name of Egyptian wheat--_Triticum vulgare_, var. _compositum_. + +Prolification of the inflorescence has been most frequently observed in +the following genera: + + _Leafy_. _Floral_. + + Ranunculaceae Ranunculus. Ranunculus! + Anemone. Anemone. + Cruciferae. *Brassica! + Caryophyllaceae. Lychnis! + Dianthus! + Geraniaceae. *Pelargonium! *Pelargonium! + Leguminosae. *Trifolium! Trifolium! + Lotus! Lotus! + Coronilla! + Cytisus. Cytisus. + Rosaceae. Poterium. + *Pyrus! *Pyrus! + *Crataegus! Crataegus! + *Rosa. Rosa! + Sanguisorba. + + Philadelphaceae. Philadelphus. + Crassulaceae. Sempervivum. + Echeveria. + Crassula. + Ficoideae. ?Tetragonia. + Cactaceae. Opuntia. Opuntia. + Pereskia. + Saxifragaceae. Saxifraga! + Umbelliferae. Seseli. + *Apium! + Cnidium. + Chaerophyllum. + Eryngium. Eryngium. + Silaus. + Heracleum! Heracleum! + Hydrocotyle. Hydrocotyle. + Daucus. + Carum. + Selinum. + Angelica! + Conium. + Astrantia. + OEnanthe. OEnanthe. + Begoniaceae. Begonia! + Valerianaceae. Valeriana. + Dipsacaceae. *Scabiosa! *Scabiosa! + Knautia! Knautia! + Compositae. *Bellis! + Centaurea. + Calendula. Calendula. + Anthemis. + Coreopsis. + Apargia. + Lampsana. + Carlina. + Arnoseris. + Tragopogon! Tragopogon! + Rudbeckia! + Senecio! + Carlina. + Bidens! Pyrethrum. + Filago. + Hedypnois. + Cirsium. + Lactuca. + Campanulaceae. Prismatocarpus. + Lobeliaceae. Jasione. + Ericaceae. Azalea! + Convolvulaceae. Convolvulus! Convolvulus! + Calystegia! + Scrophulariaceae. Scrophularia! + Antirrhinum! + Gesneraceae. Achimenes! + Primulaceae. Primula! Primula! + Cyclamen! Cyclamen! + Plumbaginaceae. Armeria. + Plantaginaceae. *Plantago! *Plantago! + Polygonaceae. Polygonum! + Euphorbiaceae. Euphorbia! + Urticaceae. Ficus. + Amentaceae. Corylus! + Castanea! Castanea. + Coniferae. *Larix! + *Cryptomeria! + Taxodium! Pinus. + Orchidaceae. Phaius! Ophrys! + Epidendrum! + Oncidium! + Liliaceae. *Allium! + *Ornithogalum! + *Lilium! + Amaryllidaceae. Fourcroya + Alismaceae. Alisma! + Palmaceae. Cocos. + Juncaceae. *Juncus! + Restiaceae. Restio! Restio! + Elegia! Elegia! + Willdenovia! Willdenovia! + Cyperaceae. Carex. + Graminaceae. Dactylis. + *Lolium! + Festuca. + *Zea! + *Triticum! + *Hordeum! + Secale. + Phleum. + +In addition to the papers already cited the following works may be +consulted with reference to prolification of the inflorescence: + + Moquin-Tandon. 'El. Ter. Veg.,' p. 376. Engelmann, 'De + Antholysi,' Secs. 85-87. Fleischer, 'Missbild. Versch. Cultur. + Pflanz.' For figures of Hen and Chicken Daisy (_Bellis + prolifera_). see Lobel, 'Ic.,' 477. Sweert, 'Florileg.,' pl. + 98, f. 5. 'Hort. Eystett. Plant. Vern.,' fol. iv, f. i. &c. For + similar malformations in marigold (_Calendula_), see Lobel, + 'Ic.,' 553. 'Act. Acad. Nat. Cur.,' vol. x, p. 208. Jaeger, + 'Missbilld.,' 192-195. 'Hort. Eystett.,' pl. aestiv. fol. iii, + f. i. Klinsmann, 'Linnaea,' t. x, p. 607. + + For monstrous plantains, in addition to previous citations, see + Camerarius, 'Epist.,' p. 261, _P. rosea_. Matthioli, + 'Krauterb,' 245. Lobel, 'Stirp. Advers. Nov.,' p. 128, _P. + major paniculata._ J. Bauhin, 'Hist. Plant.,' i, p. 503 _b_. + Ibid., p. 503, _a_, _c_, _P. major rosea_, _bracteata + paniculata_, _prolifera_, &c. 'Hort. Eystett.,' pl. aestiv., t. + vii, f. 2, _P. rosea_ et _P. bracteata_. Lobel, 'Stirp. Hist.,' + p. 162. Dodonaeus, 'Pempt.,' 1-4, cap. xxiii, P. major spica + multiplex, _i.e._ paniculata. Gerard, 'Herbal.' Clusius, + 'Plant. Rar. Hist.,' lib. v, p. 109-10, _Plantago augustifolia + Gareti prolifera_. Marchand, 'Adansonia,' iv, p. 156. + + _Coniferae._--Richard, 'Mem. Conif.,' tab. xiii, f. 9. A. Braun, + 'Das Individ.,' 1853, p. 65. De Cand., 'Organogr.,' tab. xxxvi. + Wigand, 'Bot. Untersuch.,' 154. Schlechtendal, 'Bot. Zeit.,' + 1859, p. 239. Caspary, 'De Abiet. flor. fem. struct. morphol.' + Parlatore, 'Ann. Sc. Nat.,' 1862, vol. xvi, p. 215. Cramer, + 'Bildungsabweich.,' p. 4, &c., &c. + + _Gramineae._--Bauhin, 'Pinax.,' 21. Morison, 'Hist. Plant.,' t. + i. Winckler, 'Ephem. Nat. Cur.,' dec. i, ann. 7, 8, p. 151. + Irmisch, 'Flora,' 1858, p. 40, &c. + + See also under Chloranthy, Viviparous plants, &c. + +=Prolification of the flower.=--In the preceding sections the formation +of adventitious buds of a leafy or floral nature on the inflorescence +has been considered. A similar production of buds may take place in the +flower itself, either from its centre or from the axil of some of its +constituent parts. Prolification of the flower is therefore median or +axillary, and the adventitious bud itself may be of a leafy or a floral +nature. + +=Median leafy prolification.=--In this malformation the centre of the +flower is occupied by a bud or a branch; the growing point or +termination of the axis which ordinarily ceases to grow after the +formation of the carpels, takes on new growth. This is well shown in the +accompanying illustration (fig. 58), representing the thalamus of a +strawberry prolonged beyond the fruits into a small leaf-bearing branch. + +[Illustration: FIG. 58.--Receptacle of strawberry prolonged into a leafy +branch. From the 'American Agriculturist.'] + +[Illustration: FIG. 59.--Flower of _Verbascum_ with five disunited +sepals, five similar green petals, and a prolonged branch in the centre +of the flower.] + +In other cases the carpels are entirely absent and their place is +supplied by a leafy shoot as in a species of _Verbascum_, which came +under my own observation. In this case the petals were virescent, and +the stamens and pistils were entirely absent, hence in truth, the +so-called flower more nearly resembled a branch. In a flower of a May +Duke cherry, for which I am indebted to Mr. Salter, there was a gradual +change from the floral to the foliar condition; thus there were five +distinct lanceolate sepals, the arrangement of whose veins betokened +that they were leaf-sheaths rather than perfect leaves, ten petals +partly foliaceous and sheath-like as to their venation, one of them +funnel-shaped, but whether from dilatation or cohesion of the margins +could not be determined. The stamens were eight or ten in number, their +connectives prolonged into foliaceous or petaloid appendages, so that +the filament represented the stalk of the leaf. The pistil was entirely +absent and its place was supplied by a branch with numerous perfectly +formed stipulate leaves. + +Some flowers of _Anagallis arvensis_ described by Dr. Marchand[119] are +so interesting and show so well the gradual stages by which this +malformation is arrived at, that it is desirable to cite the summary of +Dr. Marchand's researches as given in the 'Gardeners' Chronicle' by Mr. +Berkeley, taking that instance first in which the parts of the flower +departed least from the normal condition, and then the others in their +proper order. In all the parts there was a greater or less tendency to +assume a green tint; in some they were entirely green, in others the +brighter colours were confined to the more recently developed parts. + +"1. In the first case then, the sepals and petals were in their normal +position, though rather more dilated than usual; the anthers were +fertile, the principal change existing in the ovary, the upper part of +which was wanting, so that the ovules were exposed seated on the central +placenta. + +2. In the next step the calyx, more developed than usual, was separated +from the corolla by a long peduncle, and the ovary, which was ovate, +contained instead of a placenta a sort of plumule or young shoot. + +3. In this case the corolla and calyx were distant from each other; +there was no trace of stamens, but the axis was continued from the +centre of the corolla, and ended in a leaf-bud. + +4. The calyx and corolla nearly as before, but instead of stamens a +whorl of little leaves was developed, in the centre of which the axis +was continued, bearing at its tip two whorls of leaflets, alternately +three and three. + +5. In this case two out of the five stamens were normal, the other three +changed into leaves, showing clearly the origin of the leaflets, in the +last case, which took the place of the stamens. + +6. The ovary varied in different flowers. In some the placenta was +crowned with ovules; in others the ovules were replaced by a single +whorl of leaflets; in others there was every shade of change from +ordinary ovules to perfect leaflets; while in others, again, every ovule +was converted into a leaf with a long petiole. + +7. In these flowers shoots were developed in the axils of the sepals, or +on the face of the petals between the point of their insertion and that +of the stamens, and, what is most curious, in the interior of the +ovaries round the foot of the placenta. + +8. Here, again, a very singular condition presented itself: the calyx +and corolla separated from each other, the stamens partly developed, the +axis continued beyond the corolla, branched and bearing normal leaves so +as exactly to resemble an ordinary stem, while in consequence of the +calyx and corolla being bent down to the ground, adventitious roots were +developed from the axis on the under side above each of them. In another +case, where the calyx and corolla were approximated, the ovary was open +above, and sent out six shoots from within, perfectly developed, clearly +representing the central placenta and five axile buds, and each giving +out a number of adventitious roots at its base." + +In other genera of the same order (_Primulaceae_) an extension of the +placenta into a leafy branch has been observed, as in _Lysimachia_, +where in one case the prolonged placenta was removed and struck as a +cutting.[120] + +In _Ericaceae_ too, the axile placenta has been seen ovuliferous at the +base and prolonged above into a leafy branch.[121] + +=Median floral prolification.=--This is of more frequent occurrence than +the preceding. The prolonged axis is more frequently terminated by a +flower-bud than by a leaf-bud, though it must be remarked, that the +lengthened and protruded stem frequently bears leaves upon its sides, +even if it terminate in a flower, and thus the new growth partakes of a +mixed leafy and floral nature. Instances of this kind have long been +familiar to observers, and have always excited attention from the +singularity of their appearance. In one of the old stained-glass +windows, apparently of Dutch manufacture, in the Bodleian Picture +Gallery at Oxford, is a representation of a _Ranunculus_ affected with +median floral prolification.[122] In pinks the affection is not +unfrequently met with. Fig. 60 shows an instance of the kind copied from +Schotterbec. + +A singular instance of prolification in the central flower of one of the +verticillasters of _Phlomis fruticosa_ fell under my own notice; it was +a case wherein the calyx was torn on one side, and one of its lobes had +become petaloid. Between the calyx and the corolla were three or four +spathulate, hairy, bract-like organs; the corolla and stamens were +unchanged; but in place of the usual four-lobed ovary there was a single +carpel with a basilar style, terminated by a forked stigma. Occupying +the place of the other lobes of the pistil was an oblong woolly +flower-bud, consisting of calyx, corolla, and stamens, but with no trace +of pistil. I have been unable to find recorded any instance of +malformation among Labiates or Borages at all similar to this. It +differed from most other examples of prolification in that the axis was +not prolonged, the adventitious bud occupying precisely the position of +the three lobes of the ovary that were absent. The sole remaining carpel +had a style and a stigma as perfect in appearance as though the pistil +had been complete. + +[Illustration: FIG. 60.--Flower of _Dianthus_ affected with median +floral prolification.] + +In a flower of _Conostephium_ (_Epacridaceae_) forwarded to me by Mr. +Bentham, there was a similar adventitious bud placed by the side of the +pistil, but as the latter contained the usual number of cells it is +probable that the supernumerary bud in this case originated rather from +the side than the end of the axis. + +Certain families of plants present this deviation from their ordinary +structure with greater frequency than others: the following orders seem +to be the most frequently affected by it: _Ranunculaceae_, +_Caryophyllaceae_, _Rosaceae_; while it is commonly met with in +_Scrophulariaceae_, _Primulaceae_ and _Umbelliferae_. Of genera which seem +peculiarly liable to it may be mentioned the following: _Anemone_, +_Ranunculus_, _Cheiranthus_, _Dianthus_, _Dictamnus_, _Daucus_, _Rosa_, +_Geum_, _Pyrus_, _Trifolium_, _Antirrhinum_, _Digitalis_, _Primula_. + +A reference to the subjoined list of genera affected by this +malformation, and the knowledge of its comparatively greater frequency +in some than in others of them, will show that it is more often met with +in plants having an indefinite form of inflorescence than in those +having a definite one. The change may affect some only, or the whole of +the flowers constituting an inflorescence; and though it is by no means +a constant occurrence, it very frequently happens that the central or +terminal flower in a definite inflorescence is alone affected, the +others remaining in their ordinary condition, as in pinks (_Dianthus_); +and in the indefinite forms of inflorescence, it is equally common that +the uppermost flower or flowers are the most liable to be thus affected. + +In those plants which present this deviation from the ordinary condition +with the greatest frequency, it often happens that the axis is normally +more or less prolonged, either between the various whorls of the flower, +as in the case of the gynophore, &c., or into the cavity of the carpels, +as in the instances of free central placentation. To bear out this +assertion, the following instances taken from those genera having +definite inflorescence, and which are very commonly affected with +prolification, may be cited; thus, in _Anemone_ and _Ranunculus_ the +thalamus is prolonged to bear the numerous carpels; in _Dianthus_ there +is a marked internode separating the carpels from the other parts of the +flower; in _Primulaceae_ central prolification is very common, and this +is one of the orders where the placenta seems from the researches of +Duchartre and others, to be truly a production of the axis within the +carpels;[123] in _Thesium_ also, another genus with free central +placenta, this malformation has been found. + +So also among plants with indefinite inflorescence, prolification seems +very frequently to affect those wherein the axis is normally prolonged; +thus it is common in _Dictamnus_, which plant has an internode +supporting the pistil; it is frequent among _Umbelliferae_, where the +carpophore may be truly considered an axile production; it is common +among _Rosaceae_ and _Ranunculaceae_, in many of which the axis or +thalamus is well-marked, and it is by no means infrequent in the flowers +of the Orange, where the floral internodes are also slightly elongated; +on the other hand, there is no case on record in _Magnoliaceae_, and some +other orders where the floral part of the axis is at some point or other +elongated; still, on the whole, there can be but little doubt that there +is a real relation between prolification and the normal extension of the +floral internodes. + +Under these circumstances, those instances wherein the parts of the +flower become separated one from the other by the elongation of the +internodes (apostatis), constitute a lesser degree of the same change, +which operates most completely in the formation of a new bud at the +extremity of the prolonged axis. Some specimens of _Geum rivale_ (a +plant very liable to become prolified) in my possession show this very +clearly. In the wild plant the thalamus is elevated on a short stalk; in +the abnormal ones the thalamus is simply upon a longer stalk than usual, +or in a more advanced stage of the deviation the lengthened thalamus +takes the form of a branch provided with leaves and terminated by a +flower; it is noticeable, also, in these specimens, that the sepals of +the lower flower have assumed entirely the dimensions and appearance of +leaves. + +Median prolification has occasionally been recorded in flowers that +have, in their ordinary condition, but one carpel, as in _Leguminosae_ +and in _Santalaceae_. In _Leguminosae_, as also in _Amygdalus_, it would +seem as if the adventitious bud were strictly a lateral and axillary +production, and moreover that the carpel itself is not strictly terminal +but lateral in position, though apparently terminal from the abortion of +other carpels. In the only recorded instance that I am aware of, of this +malformation affecting the genus _Thesium_, the pistil was altogether +absent, and occupying its place was the new bud or branch.[124] + +[Illustration: FIG. 61.--_Daucus Carota_, showing leafly carpels, +prolification, &c.] + +As the carpels are not unfrequently absent in cases of median +prolification, it has been thought that the pistil in such cases was +metamorphosed into a stem bearing leaves or flowers. Setting aside the +physiological difficulties in the way of accepting such an opinion, an +examination of any number of cases is sufficient to refute it; for, as +Moquin well remarks, the carpels may frequently be found either in an +unaltered condition or more or less modified. + +If the pistil be normally syncarpous, its constituent carpels, if +present at all in the prolified flower, become disjoined one from the +other to allow of the passage between them of the prolonged axis; thus +in some malformed flowers of _Daucus Carota_ gathered in Switzerland +(fig. 61), not only was the calyx partially detached from the pistil, +but the carpels themselves were leaf-like, disjoined, and unprovided +with ovules; between them rose a central prolongation of the axis, which +almost immediately divided into two branches, each terminated by a small +umbel of perfect flowers, surrounded by minute bracts.[125] + +Not only are the carpels thus frequently separated one from the other by +the prolonged axis, but they undergo commonly a still further change in +becoming more or less completely foliaceous, as in the _Daucus_ just +mentioned, where the carpels were prolonged into two lance-shaped +leaves, whose margins in some cases were slightly incurved at the apex, +forcibly calling to mind the long "beaks" that some Umbelliferous genera +have terminating their fruits--for instance, _Scandix_. Dr. Norman, in +the fourth series of the 'Annales des Sciences,' vol. ix, has described +a prolification of the flower of _Anchusa ochroleuca_, in which the +pistil consisted of two leaves, situated antero-posteriorly on a long +internode, with a small terminal flower-bud between them; and numerous +similar instances might be cited. + +In this place may also be noticed those instances wherein the placenta +elongates so much that the pericarp becomes ruptured to allow of the +protrusion of the placenta, although this prolongation is not attended +by the formation of new buds. Cases of this kind occurring in +_Melastoma_ and _Solanum_ have been put on record by M. Alph. de +Candolle.[126] This is a change analogous with that which occurs in some +species of _Leontice_ or _Caulophyllum_, as commented on by Robert +Brown. See 'Miscellaneous Botanical Works' of this author, Ray Society, +vol. i, p. 359. + +If the pistil be apocarpous, and the carpels arranged spirally on an +elevated thalamus, it then frequently happens that the carpels, +especially the upper ones, become carried up with the prolonged axis, +more widely separated one from the other than below, and particularly +liable to undergo various petalloid or foliaceous changes as in +proliferous _Roses_, _Potentilla_, &c. + +[Illustration: FIG. 62.--Median floral prolification, &c., in flower of +_Delphinium_.] + +Fig. 62, copied from Cramer, shows an instance of this kind in +_Delphinium elatum_, where not only is the thalamus prolonged, and the +carpels separated, but from the axils of some of the latter which have +assumed from the disunion of their margins somewhat of the appearance of +leaves, other flowering branches proceed--axillary prolification. If, on +the other hand, the carpels be few in number, and placed in a +verticillate manner, the axis then generally passes upwards without any +change in the form or position of the carpels being apparent, as in a +proliferous columbine, figured in the 'Linnean Transactions,' vol. +xxiii, tab. 34, fig. 5. + +When a flower with the ovary naturally inferior or adherent to the calyx +becomes prolified, a change in the relative position of the calyx and +ovary almost necessarily takes place, the latter becoming superior or +detached from the calyx; this has been already alluded to in +_Umbelliferae_. In a species of _Campanula_ examined by me, the calyx was +free, the corolla double, the stamens with petaloid filaments, and in +the place of the pistil there was a bud consisting of several series of +green bracts, arranged in threes, and enclosing quite in the centre +three carpellary leaves detached from one another and the other parts of +the flower, and open along their margins, where the ovules were placed. +In other similar instances in the same species of _Campanula_, the +styles were present, forming below an imperfect tube which surrounded +the adventitious bud; in another, contrary to what occurs usually in +such cases, the ovary was present in its usual position, but surmounted +by a bud of leafy scales, enclosed within the base of a tube formed by +the union of the styles. A similar relative change in the position of +the calyx and the ovary takes place when the _Compositae_ are affected +with central prolification, or even in that lesser degree of change +which merely consists in the separation and disunion of the parts of the +flower, but which in these flowers appear to be, as it were, the first +stage towards prolification. I owe to the kindness of Professor Oliver a +sketch of a species of _Rudbeckia_? showing this detachment of the calyx +from the ovary. In a monstrous _Fuchsia_ that I have had the opportunity +of recently examining, the calyx was similarly detached from the ovary +simultaneously with the extension of the axis. Here the petals were +increased in number and variously modified, the stamens also; while in +the centre and at the top of the flower, conjoined at the base with some +imperfect stamens, was a carpel open along its ovuliferous margins. Such +instances as these seem to be the first stages of a change which, +carried out more perfectly, would result in the formation of a new bud +on the extremity of the prolonged axis. + +In _Orchidaceae_, among which family I have now met with several +instances of prolification, the ovary seems usually to be absent. Fig. +63 shows a prolified flower of _Orchis pyramidalis_ in which the +perianth was nearly regular, the central portions of the flower absent, +and their place supplied by a new miniature raceme. This specimen was +forwarded to me by Dr. Moore, of Glasnevin. + +[Illustration: FIG. 63.--Median prolification in _Orchis pyramidalis_, +the outer segments of the perianth regular and reflexed.] + +As might be expected, it very rarely happens that median prolification +occurs without some other deviation in one or more parts of the flower +being simultaneously manifested. Some of these changes have been already +mentioned, but others are commonly met with, as, for instance, the +multiplication or doubling, as it is termed, of the petals; others, +though less frequent, are of more interest. Fusion of two or more +flowers in association with prolification is especially common in +cultivated specimens of _Digitalis purpurea_; the uppermost flowers of +the raceme become fused together so as to form one large, regular, +erect, cup-shaped corolla, to the tube of which the stamens are +attached, in greater number than ordinary, and all of equal length; the +bracts and sepals are confusedly arranged on the exterior of the flower; +while in the centre, in the place usually occupied by the pistil, there +rises a conical prolongation of the axis, bearing at its outer or lower +portion a number of open carpels, provided, it may be, with styles and +ovules; these enclose an inner series of scale-like bracts, from whose +axils proceed more or less perfect florets; so that in the most highly +developed stage a perfect raceme of flowers may be seen to spring from +the centre of a cup-shaped regular flower, whose lobes show its compound +character. All intermediate stages of this malformation may be found +from cases where there is a simple fusion of two flowers with a second +verticil of carpels within the outer, up to such cases as those which +have been just mentioned. It is worthy of special remark, that in all +these cases the flowers at the uppermost part of the raceme are alone +affected, and that, in addition to the prolification, there is fusion of +two or more flowers, and regularity in the form of the compound corolla +and stamens. + +The calyx of a prolified flower is either unchanged, or it is modified +in harmony with the changes in the central part of the flower. If the +ovary be normally superior or free from the calyx, then the latter is +comparatively rarely altered; for instance, in proliferous pinks +(_Dianthus_) the calyx is seldom affected, except, indeed, in those +instances where the floral axis is prolonged, and produces from its side +a successive series of sepals, as in what is called the wheat-ear +carnation; but though these instances may be, as I believe, an imperfect +degree of prolification, they do not affect the general truth of the +above opinion, that the calyx, if it be free from the ovary, is but +rarely changed in a prolified flower; but that this is not a universal +rule is shown by proliferous flowers of _Geum rivale_, where the sepals +are usually large and leaf-like, as they likewise are frequently in +proliferous roses and pears. + +[Illustration: FIG. 64.--Proliferous rose. Hip absent, sepals leafy, +stamens wanting, axis prolonged bearing supplementary flower, &c. (Bell +Salter).] + +Proliferous roses have a special interest, inasmuch as they show very +conclusively that the so-called calyx-tube of these plants is merely a +concave and inverted thalamus, which, in prolified specimens, becomes +elongated (fig. 64) after the fashion of _Geum rivale_, &c.[127] +Occasionally from the middle of the outer surface of the urn-shaped +thalamus proceeds a perfect leaf, which could hardly be produced from +the united sepals or calyx-tube; a similar occurrence in a pear is +figured in Keith's 'Physiological Botany,' plate ix, fig. 12. + +The change which the calyx undergoes when flowers with an habitually +adherent ovary become prolified, and wherein the calyx is disjoined from +the ovary, has been before mentioned, but it may also be stated that, +under such circumstances, the constituent sepals are frequently +separated one from the other, and not rarely assume more or less of the +appearance of leaves, as in proliferous flowers of _Umbelliferae_, +_Campanulaceae_, _Compositae_, &c. + +As to the corolla, it was long since noticed that prolification was +especially liable to occur in double flowers; indeed, Dr. Hill, who +published a treatise on this subject, setting forth the method of +artificially producing prolified flowers, deemed the doubling to be an +almost necessary precursor of prolification;[128] but, though frequently +so, it is not invariably the case that the flower so affected is +double--_e.g._ _Geum_. If double, the doubling may arise from actual +multiplication of the petals, or from the substitution of petals for +stamens and pistils, according to the particular plant affected. +Occasionally in prolified flowers the parts of the corolla, like those +of the calyx, become foliaceous, and in the case of proliferous pears +fleshy and succulent. There is in cultivation a kind of _Cheiranthus_? +in which there is a constant repetition of the calyx and corolla, +conjoined with an entire absence of the stamens and pistils; a short +internode separates each flower from the one above it, and thus +frequently ten or a dozen of these imperfect flowers may be seen on the +end of a flower-stalk, giving an appearance as if they were strung like +beads, at regular intervals, on a common stalk. I have seen a similar +instance in a less degree in a species of _Helianthemum_. + +The stamens are subject to various changes in prolified flowers; they +assume, for instance, a leaf-like or petal-like condition, or take on +them more or less of a carpellary form, or they may be entirely absent; +but none of these changes seem to be at all necessarily connected with +the proliferous state of the flower. Of more interest is the alteration +in the position of these organs which sometimes necessarily accrues from +the elongation of the axis and the disjunction of the calyx; thus, in +proliferous roses the stamens become strictly hypogynous, instead of +remaining perigynous. In _Umbelliferae_ the epigynous condition is +changed for the perigynous, &c. + +The condition of the pistillary organs in prolified flowers has already +been alluded to. Hitherto those instances have been considered in which +either the carpels were absent, or the new bud proceeded from between +the carpels. There is also an interesting class of cases where the +prolification is strictly intra-carpellary; the axis is so slightly +prolonged that it does not protrude beyond the carpels, does not +separate them in any way, but is wholly enclosed within their cavity. +Doubtless, in many cases, this is merely a less perfect development of +that change in which the axis protrudes beyond the carpels. This +intra-carpellary prolification occurs most frequently in plants having a +free central placenta, though it is not confined to them, as it is +recorded among _Boragineae_. A remarkable instance of this is described +by Mr. H. C. Watson in the first volume of Henfrey's 'Botanical +Gazette,' p. 88. In this specimen a raceme of small flowers was included +within the enlarged pericarp of a species of _Anchusa_. But the most +curious instances of this form of prolification are, no doubt, those +which are met with among _Primulaceae_ and other orders with free central +placentation. + +Duchartre, in his memoir on the organogeny of plants with a free central +placenta, in the 'Ann. des Sc. Nat.,' 3 ser., 1844, p. 290, among other +similar instances, mentions two flowers of _Cortusa Matthioli_, wherein +the placenta was ovuliferous at the base; but the upper portion, instead +of simply elongating itself into a sterile cone, had produced a little +flower with its parts slightly different from those of the normal +flowers. M. Alph. de Candolle has likewise described somewhat similar +deviations, and one in particular in _Primula Auricula_, where the +elongated placenta gave off long and dilated funiculi bearing ovules, +while other funiculi were destitute of these bodies, but were much +dilated and foliaceous in appearance.[129] In some flowers of +_Rhododendron_ I have observed a similar condition of the ovules, which, +moreover, in the primary flowers, were attached to the walls of the +carpels--parietal placentation. + +In speaking of these as cases of intra-carpellary prolification, it is, +of course, impossible to overlook the fact that they differ in degree +only from those cases where the lengthened axis projects beyond the +cavity of the carpels; nevertheless they seem to demand special notice, +because in these particular plants the placenta or its prolongation +appears never to protrude beyond the carpels, or at least very rarely. +There are, however, numerous instances of such an extension of the +placenta and of prolification occurring among _Primulaceae_ in +conjunction with the more or less complete arrest of growth of the +carpels.[130] An instance of this kind has come under my own notice in a +monstrosity of the chinese primrose, in which the carpels were reduced +to a hardly discernible rim surrounding an umbel of five rays, each +terminated by a small normally constituted flower-bud. + +The ovules of a prolified flower are either unaffected, or they occur in +a rudimentary form, or, lastly, they may be present in the guise of +small leaves. + +Under the term prolification of the fruit two or three distinct kinds of +malformation appear to have been included. The term seems usually to be +applied to those cases where from the centre of one fruit a branch +bearing leaves, flowers, or another fruit, is seen to project, as +happens occasionally in pears. Now, in many instances, not only the +fruit, is repeated, but also the outer portions of the flower, which +wither and fall away as the adventitious fruit ripens; so that at length +the phenomenon of one fruit projecting from another is produced. It is +obvious that this form of prolification in no wise differs from ordinary +central prolification. Sometimes some of the whorls of the adventitious +flower are suppressed; thus, M. Duchartre describes some orange blossoms +as presenting alternating series of stamens and pistils one above +another, while the calyces and corollas belonging to each series of +stamens and pistils were entirely suppressed.[131] In other cases, +doubtless, the carpellary whorl is alone repeated, the other whorls of +the adventitious flower being completely absent. + +Another condition, apparently sometimes mistaken for prolification of +the fruit, is that in which the carpellary whorl becomes multiplied; so +that there is a second or even a third series within the outer whorl of +carpels. If the axis be at all prolonged, then these whorls are +separated one from the other, and produce in this way an appearance of +prolification. This happens frequently in oranges, as in the variety +called Mellarose.[132] + +Moquin has given an explanation of the St. Valery Apples, wherein the +petals are sepaloid, the stamens absent, and where there is a double row +of carpels, by supposing these peculiarities to be due to "a +prolification combined with penetration and fusion of two or more +flowers," but it is surely more reasonable to conceive a second row of +carpels placed above the first by the prolongation of the central part +of the axis. Supposing this view to be correct, the inner calyx-like +whorl might be considered either as a repetition of the calycine whorl, +or it might be inferred that the corolla was present in the guise of a +second calyx. + +Moquin-Tandon suggests another explanation--namely, that though the +stamens are absent in these curious flowers, at least in their ordinary +shape, they are represented by the lower row of carpels, which become, +in process of development, fused with the upper or true carpels. If this +were so, surely some intermediate conditions between stamen and carpel +would occasionally be present; but such does not appear to be the +case.[133] + +In some of the instances of so-called proliferous pears the carpels +would seem to be entirely absent, and the dilated portion of the axis to +be alone repeated. Thus, the axis dilates to form the lower fruit +without any true carpels being produced, but at its summit a whorl of +leaves (sepals) is formed; above these another swelling of the axis +takes place also without the formation of carpels, and this, it may be, +is terminated in its turn by a branch producing leaves. In these cases +there is no true prolification, but simply an extension of the axis. +That the outer portion (so-called calyx-tube) of these fruits is really +an axile product there can now be little doubt; and, as if to show their +axile nature, they occasionally produce leaves from their sides, as +before mentioned. Moquin, in the tenth volume of the 'Bulletin of the +Botanical Society of France,' p. 73, says that when the case is one of +prolification the lower fruit is larger and is formed of a fleshy mass; +moreover, the line of demarcation between the fruits is more distinct, +and there are traces of the seed-bearing cavity in the interior, and of +calycine lobes at the top. On the other hand, if the case be one of +hypertrophy merely, the lowermost fruit is the smallest, and there is no +trace of seed-bearing cavity nor of sepals. See also under Hypertrophy. + +Some other malformations usually referred to prolification of the fruit +seem due to branching of the inflorescence, as in _Plantago_, wheat, +maize; or to a simple extension of the axis beyond its ordinary limit, +as in some cones of firs, &c. It is obvious that the true fruits in +these cases are in no wise affected. + +From these considerations it would appear better to abandon the use of +the expression prolification of the fruit, as unnecessary where it is +really applicable, and as delusive in the numerous other cases where it +is employed. + +Median prolification of one or other kind has been met with in the +following genera: + + _Leafy_. _Floral_. + + Ranunculaceae. Clematis. + Anemone! *Anemone! + Ranunculus! *Ranunculus! + Delphinium. + Caltha. + Aquilegia! + Cruciferae. Bunias. + *Cheiranthus! + Erucago. + *Matthiola! + Sisymbrium! + Brassica! + Nasturtium. + Hesperis. + Sinapis! + Diplotaxis. + Lunaria. + Erysimum. + Alyssum. + Peltaria. + Cardamine! + Cleome. + Cistaceae. Helianthemum! + Caryophylleae. Dianthus! *Dianthus! + Silene! + Lychnis! + Violaceae. Viola! + Tiliaceae. Triumfetta! + Geraniaceae. Geranium! + Sapindaceae. Pavia! Pavia! + Malvaceae. Paritium. + Hibiscus! + Malpighiaceae. Byrsonima! + Rutaceae. Genera not specified. *Dictamnus! + Resedaceae. Reseda. + Caylussa! + Aurantiaceae. *Citrus! + Vitaceae. Vitis. Vitis. + Umbelliferae. Heracleum. + Angelica. + Thysselinum. + *Athamanta. + *Daucus! + *Torilis. + Rosaceae. *Rosa! *Rosa! + *Geum! *Geum! + Agrimonia. Amygdalus. + Prunus! + Spiraea! Spiraea! + Rubus. + *Pyrus! *Pyrus! + ?Leguminosae Trifolium! + Medicago! + Melilotus. + Pisum! + Cucurbitaceae. Cucumis. + Passifloraceae. Passiflora. + Philadelphaceae. Philadelphus. + Onagraceae. Epilobium! + Epacridaceae. Epacris! + Ericaceae. *Erica. + Rhododendron! + Convolvulaceae. Convolvulus. + Gentianaceae. Gentiana. Gentiana. + Apocynaceae. Vinca. + Jasminaceae. Jasminum! + Scrophulariaceae. Verbascum! Antirrhinum! + *Digitalis! + *Linaria! + Veronica. + Orobanchaceae. Orobanche. + Labiatae. Genera not specified. Stachys. + Phlomis! + Hydrophyllaceae. Hydrophyllum. + Boraginaceae. Anchusa. + Symphytum. + Primulaceae. *Dodecatheon. *Cortusa. + *Anagallis! *Anagallis! + *Primula. + Dipsacaceae. Scabiosa. + Compositae. Hieracium! Hieracium! + Cirsium. Cirsium. + Hypochaeris. Calendula! + Spilanthes. + Carthamus. + Coreopsis. + Campanulaceae. Campanula. *Campanula! + Polygonaceae. Genera not specified. Rumex. + Santalaceae. Thesium. + Liliaceae. Genera not specified. Tulipa! + Hemerocallis! + Asphodelus. + Hyacinthus! + Iridaceae. Iris. + Amaryllidaceae. Narcissus! + Leucojum. + Orchidaceae. Orchis! + Habenaria. + Cyperaceae. Carex. + Gramineae. Phleum. + +=Axillary prolification= is the term applied to those cases wherein one +or more adventitious buds spring from the axils of one or more of the +parts of the flower. Engelmann makes use of the word ecblastesis to +denote the same condition. Both terms are open to the objection that +they do not clearly enable us to distinguish prolification occurring +within the flower from a similar state originating outside the flower, +within the bracts of the inflorescence. This latter condition, called by +Moquin-Tandon lateral prolification (see Prolification of the +Inflorescence), is as truly axillary as that to which the name is +restricted. In consequence of certain peculiarities in the structure of +some flowers, to be hereafter alluded to, it is not in all cases easy to +decide whether the new growth springs from the interior of the flower, +or from the inflorescence beneath the flower. + +The accessory bud presents itself as a leaf-bud, a branch, a flower-bud, +or a miniature inflorescence; it may be sessile, but is far more +frequently stalked, and in more than half the number of cases it is a +flower-bud or an inflorescence. There may be one or more of these buds; +if two only, then they are usually placed directly opposite one to the +other, on the opposite sides of the flower. + +It will be seen, from the appended list, that the orders and genera in +which this description of adventitious growth occurs most frequently are +the following:--_Cruciferae_, especially the genus _Brassica_; +_Caryophyllaceae_, e.g. _Dianthus_; _Resedaceae_; _Leguminosae_, e.g. +_Melilotus_, _Trifolium_, &c.; _Rosaceae_, e.g. _Rosa_, _Potentilla_, +&c.; _Umbelliferae_, and _Campanulaceae_. For the most part, these are +groups also peculiarly liable to central prolification. + +All the parts of the flower may be thus affected; but, as might have +been anticipated from the foliaceous nature of the sepals, the new bud +usually arises from within the axil of one of those organs. Next in +frequency to the calyx, the pistil is subjected to this change--the +carpels in such a case being disunited and leaf-like. The petals rank +next, and lastly the stamens; these latter, indeed, are usually, but not +invariably, absent, the new growth occupying their position. Hence it +may well be that when such is the case, there is no real axillary +prolification, but rather the substitution of a bud for a stamen. +Generally, however, the position of the accessory bud is such that it +may properly be referred to the axil of an undeveloped or rudimentary +stamen. + +The largest number of instances of this malformation, not merely +generically, but also individually, occurs in plants the members of +whose floral whorls are not united one to the other; thus, it is far +more common in polypetalous plants than in gamopetalous ones. In the +prolified flowers belonging to the latter group, the sepals, if not +actually uncombined, are only united for a short distance. The same +relationship, but in a much less degree, exists in the case of median +prolification, as that aberration is likewise most commonly met with in +polypetalous flowers. Another feature of interest is the rarity with +which axillary prolification is found in irregular gamopetalous blooms. +It may be that the irregular and comparatively excessive growth in some +parts of these flowers, as compared with others, may operate in checking +any luxuriant tendency in other directions. + +As in the case of median prolification, plants having an indefinite +inflorescence are more liable to be affected with ecblastesis than those +having a definite one. The degree of branching of the inflorescence may +be noticed, as this deformity is far more common in plants whose +peduncles are branched than in those which have either a solitary flower +or an unbranched flower-stalk. More than two thirds of the entire number +of genera cited as the subjects of this malformation have a branched +inflorescence of some form or other; and about two thirds of the cases +occur in genera having some form of indefinite inflorescence. If +individual instances could be accurately computed, the proportion would +be even higher. + +Fully three fourths of the entire number of genera recorded as +occasionally the subjects of this irregularity possess in their usual +state some peculiarity of the thalamus; for instance, in about a third +of the whole number of genera the thalamus is more or less prolonged +between some or other of the floral whorl, e.g. _Caryophyllaceae_, +_Potentilla_, _Anemone_, _Dictamnus_, _Umbelliferae_, &c. About one +fourth of the genera have numerous stamens or numerous carpels, or both, +springing naturally from the thalamus. In others (about one sixth) the +thalamus is enlarged into a disc, or else presents one or more +glandular swellings, _e.g._ _Reseda_, _Nymphaea_, _Cruciferae_. In the +last-named family, as has been already remarked, prolification is very +common. It would be interesting to ascertain precisely what part of an +inflorescence is most liable to this affection; but as information on +this point is but rarely given in the records of these cases, I can only +give the results of my own observations, which go to show that, in a +many-flowered inflorescence, those flowers at the outside, or at the +lower portion, seem to be more frequently the subjects of this change +than those situated elsewhere. This may probably be accounted for by the +fact that the malformation is met with most generally in plants with an +indefinite form of inflorescence, and therefore the lowermost or +outermost flowers are most fully nourished; the upper flowers being in a +less advanced condition, the change is more likely to be overlooked in +them; or it may be that from the unusual luxuriance in the lower +flowers, the upper ones may be either present in their ordinary +condition, or may be (as indeed frequently happens) stunted in the size +and proportion of their several parts. + +=Axillary foliar prolification of the flower.=--The formation of an +adventitious leaf-bud in the axil of any of the parts of the flower is +not of such common occurrence as the development of a flower-bud in +similar situations, nor is it so frequent as median foliar +prolification. I have seen leafy shoots proceeding from the axils of the +sepals in the flowers of _Brassica_, and a similar occurrence has been +noticed in _Caltha palustris_, _Herreria parviflora_, and other plants. +Dr. Marchand's flowers of _Anagallis_, previously referred to at p. 117, +showed good illustrations of this occurrence, as also some specimens +described by Kirschleger in _A. phoenicea_.[134] Steinheil has figured +and described[135] a flower of _Scabiosa_ in which there was an +adventitious formation of leafy shoots in the axil of the outer calyx. +In some flowers, such as _Convolvulus_, _Anemone_, &c., the exact nature +of the sub-floral leaves is uncertain, _i.e._ it is open to doubt +whether the organs in question are bracts or leaves pertaining to the +inflorescence, or whether they are really parts of the flower. When +leafy shoots are formed in the axils of such organs, the adventitious +growth may be referred to extra-floral prolification, prolification of +the inflorescence that is, or to axillary prolification, according to +the view taken of the real nature of the sub-floral leaves. So far as +the mere occurrence of prolification is concerned, it is not very +material which view be adopted. The same remark applies to cases where +leaf-buds occur on the outer surface of inferior ovaries, as in +_Rosaceae_, _Pomaceae_, _Philadelphus_, or _Tetragonia expansa_, as +elsewhere mentioned. + +It would seem more consistent with the general arrangements of parts, +that the adventitious buds should be formed more frequently outside than +within the flower proper. + +Knight[136] figures and describes the occurrence of small tubers or +fleshy leaf-buds in the axils of the sepals of a potato, a curious +illustration of the real morphological nature of the tuber. + +=Axillary floral prolification of the flower.=--As already stated, this +is of more common occurrence than the formation of a leaf-bud in a +similar situation. Any of the parts of the flower may thus subtend a +flower-bud, though probably the new buds more frequently originate in +the axils of the sepals than in the other whorls. In _Cruciferae_ the +change in question is, relatively speaking, very common. In cauliflowers +and broccoli I have frequently met with stalked flowers proceeding from +the axils of the sepals, so also in some fuchsias I have seen a ring of +stalked flower-buds alternating with the petals, which, together with +the stamens and pistil, remained unaffected. The number of parts in the +supernumerary structures is generally less than the normal flowers. + +In Mr. Herbert Spencer's 'Principles of Biology,' part iv, p. 37, are +figured and described some monstrous inflorescences in _Angelica_ and +other _Umbelliferae_, from which, amongst other things, the author draws +the conclusion that there is no absolute distinction between leaf and +branch. Without staying for the moment to discuss this matter, it may +here be said that the Umbellifers in question apparently owe their +peculiarities rather to axillary prolification within the flower, or to +prolification of the inflorescence, than to an actual transformation of +a flower or any portion of a flower into an umbellule.[137] + +In the 'Gardeners' Chronicle,' 1855, p. 551, an instance is figured of +the production of a supernumerary flower proceeding from the axil of a +stamen in a species of _Nymphaea_ (fig. 65). The ovary in this case was +wanting, but in its place was a tuft of small leaves. It is curious that +among Dr. Kirk's drawings of east tropical African plants now at Kew, +there should be one representing a precisely similar state of things. +The species in both instances was _Nymphaea Lotus_, or a cultivated +variety of it. + +M. Wesmael[138] describes a very singular case of what appears to have +been referable to axillary prolification in the flowers of _Carex +acuta_. The rachillus is described as prolonged through the utricle by +the side of the stigmas, bearing on its side a bract, then a secondary +utricle, from the axil of which sprung a short stem surmounted by an +ovary. Wigand, 'Flora,' 1856, mentions a similar change in _Carex +glauca_. In this instance the base of the female inflorescence bore +lateral spikes, which projected from the utricles; some of these +adventitious spikes were female, others female below and male above, +others, again, wholly male. + +[Illustration: FIG. 65.--Flower of _Nymphaea Lotus_, var., showing +axillary floral prolification. The section also shows the tuft of leaves +that occupied the place of the ovary.] + +Various changes in the form and arrangement of the several floral whorls +accompany axillary prolification; some of these affect the particular +organ or organs implicated, and these only, while in other cases some +other parts of the flower likewise undergo modification. The changes +most commonly met with are such as may be classed under Goethe's theory +of retrograde metamorphosis; for instance, if a supplementary bud be +developed in the axil of a sepal, that sepal is likely to be more than +ordinarily leaf-like in appearance. The dislocation of the affected +sepal from its fellows is a very frequent occurrence; in cases of this +kind the detached sepal is placed below the others, thus approximating, +in position as well as in function, to the bracts. In some of the +instances of proliferous pears, on which I shall have occasion to +comment, the sepals are described as sharing in the succulent character +of the fruit. + +The petals, under such circumstances, often exist in the guise of sepals +or of small leaves; and instances are recorded wherein the place of the +calyx and corolla was supplied by a succession of overlapping green +scales, from the axils of which the new buds arose. M. Germain de Saint +Pierre records such a case in _Trifolium repens_, wherein the calyx and +corolla were replaced by overlapping scales, in the axils of each one of +which arose a flower; above there was a row of stamens, and in the +centre a pistil in the guise of a trifoliate leaf.[139] Such instances +seem to afford an extreme degree of a more common change, viz., the +diminished size and contracted appearance of the sepals and petals when +affected with axillary prolification. They have also a close +relationship to such developments as we see in the wheat-ear carnation, +in certain species of the genus _Maesa_ and others, wherein the calyx is +repeated over and again, to the partial or complete suppression of the +other parts of the flower. All these cases may be in part explained by +the operation of the principle of compensation. + +So far as the androecium is concerned, the stamens either remain +unaltered, or they are present in a more or less petal-like condition; +but it far more frequently happens that the stamens are entirely +suppressed, the adventitious bud supplying their place; thus was it in +the _Dianthus_ represented in the adjoining woodcut, fig. 66, where the +stamens were entirely absent, and their places supplied by +flower-bearing branches. This _Dianthus_ has the more interest from its +similarity to the one described by Goethe, Metam. der Pflanzen, cap. +16, sect. 105; but in that instance median prolification also existed. +For my specimens I am indebted to Mr. T. Moore. + +[Illustration: FIG. 66.--Flower of _Dianthus_ sp., calyx removed; petals +turned down so as to show the stalked flower-buds springing from their +axils.] + +The pistil, too, is necessarily subject to very grave alterations when +affected with this malformation. It is separated into its constituent +carpels; and these assume a leaf-like aspect, and are in the great +majority of instances destitute of ovules. Indeed, virescence or +chloranthy is very intimately connected with this aberration, as might +have been anticipated, for if the parts of the flower assume more or +less of the condition of stem-leaves or bracts, it is quite natural to +expect that they will partake likewise of the attributes of leaves, even +at the expense of their own peculiar functions. + +It occasionally happens that an adventitious bud arises from the axil of +a monocarpellary pistil. This takes place sometimes in _Leguminosae_, and +seems to have been more frequently met with in _Trifolium repens_ than +in other plants. The species named is, as is well known, particularly +subject to a reversion of the outer whorls of the flower to leaves, and +even to a leaf-like condition of the pistil. There are on record +instances wherein a leaf-bud has been placed in the axil of a more or +less leaf-like carpel; while at other times a second imperfect carpel +has been met with in the axil of the first.[140] I have myself seen +numerous imperfectly developed cases of this kind. + +It may be asked whether such cases are not more properly referable to +central prolification--whether the axis is not in such flowers +terminated by two, rather than by one carpel? It is, however, generally +admitted by morphologists that the solitary carpel of _Leguminosae_ is +not terminal, but is the sole existing member of a whorl of carpels, all +the other members of which are suppressed as a general rule, though +exceptional instances of the presence of two and even of five carpels +have been described.[141] + +Again, the adventitious bud or carpel is placed, not laterally to the +primary one, or opposite to it, on the same level, but slightly higher +up--in fact, in the axil of the primary carpellary leaf. Griffith +figures and describes[142] an instance of the kind in a species of +_Melilotus_. The stalk of the ovary is mentioned as having a sheathing +base, bearing in its axil a prolongation of the axis of inflorescence, +in the form of a short spike with hairy bracts and imperfect flowers, +the latter having a well-formed calyx and rudimentary petals and +stamens. Griffith infers, from this specimen, that the legume is not to +be considered as a terminal leaf. + +_List of Genera in which Axillary Prolification has been observed._ + + Order Genus. Leaf-bud Flower-bud or From what organ. + or Branch Inflorescence + + Ranunculaceae Clematis Flower-bud Sepals. + Caltha Ditto Ditto. + Aconitum Ditto. + Delphinium Ditto Sepals, carpels, + &c. + Anemone! Ditto Involucre? + Nymphaeaceae Nymphaea! Fruit? + Nymphaea Flower Petal. + Cruciferae *Brassica! Leaf-bud Flower-bud Sepals and + petals. + Brassica! Ditto Stamens. + Brassica! Ditto Ditto Pistil. + Cardamine! Ditto Sepals. + Matthiola! Ditto Sepals and + petals. + Cheiranthus! Ditto Sepals. + Erysimum Ditto Sepals and + pistils. + Lepidium! Ditto Petals and + stamens. + Arabis Ditto Sepals. + Diplotaxis Flower, Pistil, calyx + inflorescence and corolla. + Capsella + Capparidaceae Cleome Flower-bud Sepals. + Resedaceae *Reseda Ditto Ditto. + Caryophyllaceae Arenaria Branch Ditto. + Agrostemma Leaf-bud Ditto. + *Lychnis Ditto + Stellaria Ditto + Silene Ditto + *Gypsophila Ditto Ditto Sepals and + stamens. + *Dianthus! Ditto Ditto Sepals. + Dianthus! Ditto Inflorescence Petals and + stamens. + Cucubalus Sepals + Saponaria! Sepals and + petals. + Malvaceae Alcea Flower-bud Stamen. + Aurantiaceae Citrus! Ditto Ditto. + Rutaceae Dictamnus! Ditto Pistil leafy. + Tropaeolaceae Tropaeolum! Ditto Petals. + Celastraceae Celastrus Ditto Sepals. + Leguminosae *Melilotus! Inflorescence Sepals and + petals. + Medicago Flower-bud Sepals. + Coronilla Ditto Ditto. + Trifolium! Ditto Second carpel Pistil. + axillary to + first + Melilotus! Ditto Ditto + Trifolium! Flower-bud Sepals and + petals. + Rosaceae Pyrus! Fruit? Fruit? + Cerasus! Flower-bud Petals and + stamens. + Potentilla! Ditto Leafy carpels. + Crataegus! Ditto Petals. + *Rosa! Ditto Ditto Sepals, petals, + stamens and + pistil. + Myrtaceae Lecythis Ditto Fruit? + Tetragoniaceae Tetragonia? Ditto Ditto. + Cactaceae Opuntia! Fruit-like Tufts of spines. + branch + Pereskia Ditto Sepals? + Echinocactus Ditto Ditto. + Philadelphaceae Philadelphus Ditto Sepals. + Umbelliferae *Athamanta Ditto Calyx. + *Daucus! Ditto Calyx and + pistil. + Bupleurum Ditto Calyx and + pistil. + Torilis Ditto Calyx and + pistil. + Apium Flower-bud Calyx and + pistil. + Pastinaca Ditto Ditto ditto. + Heracleum! Ditto Ditto ditto. + Angelica! Umbel Ditto ditto. + Campanulaceae *Campanula! Branch Sepals. + Prismatocarpus Ditto Fruit Sepals, &c. + Gentianaceae Gentiana! Flower-bud Sepals. + Convolvulaceae *Convolvulus! Ditto Outer calyx. + Solanaceae Solanum! Ditto Sepals. + Solanum Tubers Sepals and + petals. + Scrophulariaceae *Digitalis! Ditto Petals, &c. + Veronica Raceme Calyx. + Primulaceae Anagallis! Branch Ditto Petals. + Primula Ditto Petals and + carpels. + Polygonaceae Rumex Ditto Sepals. + Santalaceae Thesium Leaf-bud In place of + stamens and + pistils, both + absent. + Euphorbiaceae? Euphorbia? Ditto Outer bracts? + Orchidaceae Orchis! Flower-bud Perianth. + Amaryllidaceae Leucoium Ditto Ditto. + Iridaceae Iris Ditto Pistil. + Liliaceae Herreria Ditto Sepals. + Hyacinthus Flower and Perianth. + raceme + Convallaria Flower-bud Ditto. + Allium Ditto Ditto. + Cyperaceae Carex Inflorescence Utricle. + +[Illustration: FIG. 67.--Proliferous Rose. Calyx leafy; petals normal, +some reflexed; stamens and pistil absent; in their places a branch with +leaves and flowers.] + +[Illustration: FIG. 68.--Rose exhibiting median, axillary, lateral, +floral, and leafy prolification in same flower.] + +=Complicated prolification.=--From what has been before stated it may be +seen that prolification of two or more kinds may coexist in the same +flower. Mixed leafy and floral prolification is not unfrequent in +proliferous roses, where a shoot is, as it were, prolonged through the +centre of the original flower and terminated by a second flower, or +even by a cluster, as is well shown in the accompanying figure (fig. +67). Median and axillary prolification, also, not unfrequently coexist +in the same flower; thus, in a proliferous rose forwarded to me by Mr. +W. Thomson (fig. 68), the following changes were observed:--the swollen +portion below the calyx, the "hip," was entirely absent; the sepals were +leaf-like in aspect, the petals unaffected; above the petals the axis +was prolonged for a short distance and then bore a circlet of miniature, +sessile roses, destitute, indeed, of calyx, but provided with numerous +petals, stamens, and pistils. Above these lateral flowers, the prolonged +axis bore a number of scales in many rows. The scales were in their turn +surmounted by a whorl of five perfect leaves, beyond which, again, the +axis was prolonged into a leafy shoot terminated by a flower bud, the +whole constituting a remarkably complicated admixture of elements +belonging to the flower, the bud, the inflorescence, and the +leafshoot.[143] + +Proliferous flowers of Orchids also occasionally present great +complexity in the arrangement of their parts. An instance of this kind +was described by myself from specimens furnished by Dr. Moore, of +Glasnevin, in the 'Journal of the Linnean Society,' vol. ix, p. 349, +tabs. x, xi, and from which the following summary is extracted: + +[Illustration: FIG. 69.--Proliferous Orchis. Diagram showing the +arrangement of the several organs in the seven outer circles of the +flower. Each whorl is numbered, and the position of the axillary buds +shown by the small circles.] + +The primary flowers were composed of five distinct whorls, and of at +least two others less perfectly developed. These primary flowers did not +give rise to median formations, but they produced secondary buds in the +axils of the segments of the perianth. These latter buds were themselves +the subject of tertiary prolification of both kinds, median and +axillary. The tertiary median growths, like the primary flower, did not +develop median buds, but only lateral ones--quaternary axillary +prolification. + +The accompanying diagrams are intended to show the plan of arrangement +in these flowers. Fig. 69 shows the disposition of parts in the primary +flower and the situation of the axillary buds. Fig. 70 shows the primary +flower without any central prolongation, but giving off axillary buds, +two of which are shown in the diagram, 2, 2; these are, each of them, +the subject of both median, 3, 3, and axillary prolification, 4', 4'. + +[Illustration: FIG. 70.--Diagram to explain the construction of the +double-flowered _Orchis_. + +1. The primary flower, with no median bud, the position of which, had it +been present, is shown by the dotted line. + +2. Two axillary buds proceeding from 1, and themselves giving origin to + +3, 3. Median buds, and 3', 3', axillary buds. + +4' 4'. Axillary buds, proceeding from 3. No median bud is produced from +3; its situation, had it been present, is indicated by the dotted line.] + +In _Narcissus major_ a similar combination of both forms of +prolification exists, as described by Morren.[144] + +On the general subject of Prolification in flowers, in addition to the +authorities already cited, the reader may refer to the following among +many others: + + Linnaeus, 'Prolepsis,' Secs. vi et vii. Goethe, 'Versuch. + Metamorph.,' cap. xv and xvi Secs. 103-106. Moquin-Tandon,' El. + Ter. Veg.,' p. 362, &c. Engelmann, 'De Antholys.,' Secs. 52-62, + &c. Cramer, 'Bildungsabweichungen,' &c. _Orchidaceae_, + _Umbelliferae_, _Compositae_, _Leguminosae_, _Primulaceae_, + _Ranunculaceae_. Fleischer, 'Missbild. Cultur Gewachs.' + Schlechtendal, 'Linnaea,' xv, p. 408, _Rosa_. 'Bot. Zeit.' vol. + xx, 1862, p. 382, _Cyclamen_. 'Bot. Zeit.,' vol. xx, p. 301, + _Asphodelus_; et _Lilium_. Seringe, 'Bull. Bot.,' i, t. xi, f. + 7, 8, _Arabis_, _Diplotaxis_. Clos, 'Mem. Acad. Toulouse,' 5th + ser., 1862, _Papaver_. Wigand, 'Flora,' 1856, p. 716, + _Hypochaeris_; et 'Bot. Untersuch.,' p. 19. Buchenau,' + Flora,' 1857, p. 295, _Reseda_. Roeper, 'Bot. Zeit.,' 1852, p. + 427, _Orchis_. Presl., 'Linnaea,' vi, p. 599, tab. ix, figs. + 5-8, _Sisymbrium_, Vrolik., 'Flora,' 1846, p. 97, t. i et ii, + id. 1844, t. i, _Digitalis_. See also Schlechtendal, 'Bot. + Zeit.,' vol. ix, 1851, p. 579. Klinsmann, 'Linnaea,' x, p. 604, + t. v, _Hesperis_. Fuckel, 'Flora,' 1848, p. 609. _Melilotus_. + De Candolle, 'Organogr.,' i, 396, t. 33. Turpin, 'Atlas de + Goethe,' p. 65, t. 5, figs. 12, 13. Fenzl. 'Sitzungsbericht d. + k. Akad. d. Wissensch. Wien.,' heft, iii, tabs. 3, 4, _Rosa_. + Kirschleger, 'Flora,' 1845, 613, _Dianthus_, _Rosa_. + 'Institut.,' 1841, No. 413, p. 421, _Tragopogon_. Baron de + Melicoq., 'Ann. Sc. Nat.,' 3rd ser., vol. v. 1846, p. 61, + _Antirrhinum_. Reichenbach, 'Icon. Fl. Germ.,' tab. 100, + _Reseda_--"monstrosa anticipatio Euphorbiacearum et + Capparidearum." Duhamel, 'Phys. Arbres.,' liv. iii, cap. 3, p. + 303, pl. xii, f. 306, _Rosa_. Caspary, 'Bull. Soc. Bot. Fr.,' + vol. vi, 1859, p. 235, Rev. Bibl., _Pyrus_. Eichler, 'Flora,' + 1865, tab. ix, _Cleome_. Lindley, 'Elements of Botany,' p. 63, + &c., _Rosa_, _Epacris_, _Anagallis_, _Pyrus_. Irmish, 'Flora,' + 1858, p. 38, _Pyrus_; and 'Bot. Zeit.,' xix, 1861, p. 342, + _Hyacinthus_. Duchartre, 'Bull. Soc. Bot. France,' 1861, p. + 451, _Rosa_. Weber, 'Verhandl. Nat. Hist. Verein. Rhein. + Preuss., &c.' 1858 et 1860. Landrin, 'Mem. Soc. Sc. Nat. Seine + et Oise,' 1866?[145] Masters, 'Trans. Linn. Soc.,' vol. xxiii, + p. 359, tab. 34 and p. 481, tab. 54. + +=Prolification of the embryo.=--This term was applied by Moquin-Tandon +to a peculiar condition of the almond (_Amygdalus_), in which, indeed, +it is not of unfrequent occurrence. In these cases one almond encloses +within its cotyledons a second embryo, and this, again, in some +instances, a third, the little plants being thus packed like so many +boxes one within the other. The supplementary embryos are, in the ripe +state at least, quite separate and detached one from another. These +cases differ from the ordinary instances wherein there is an increased +number of embryos in one seed in their position. In the latter case, as +often happens in the seeds of the orange, the new products are placed by +the side one of another.[146] + +For other cases of prolification or the adventitious formation of buds +on leaves, roots, &c., see under Heterotaxy. + +FOOTNOTES: + +[104] Linn., 'Prolepsis,' Sec. vii; Goethe, 'Metamorph.,' Secs. 96, +103, 106. + +[105] "Diaphysis inflorescentiarum." Engelmann, 'De Anthol.,' Sec. 85. + +[106] 'Flora,' 1844, p. 565. + +[107] 'Ann. Sc. Nat.,' ser. 3, vol. v, 1846, p. 64. + +[108] 'Bot Zeit.,' vol. xx, p. 382. + +[109] 'Miscel. Austriac. Bot.,' vol. i, Vindob, 1778, p. 133. + +[110] "_Umbellati dum prolificantur, augent umbellulam, ut ex umbellula +simplici altera exeat_." 'Linn. Phil. Bot.,' Sec. 124. + +[111] 'En. Euphorb.,' p. 36. + +[112] Meisner. 'Mon. Gen. _Polygoni_ Prodrom.,' p. 20, tab. v, considers +the bulbils of this plant to be modifications of the pedicels of the +flower. + +[113] See A. Braun. 'Ann. Scienc. Nat.,' 4th series, 1860, vol. xiv, p. +13. + +[114] "_Prolificatio e latere ex calyci communi proles plurimos +pedunculatos emittens, fit in compositis aggregatis proprie dictis._" +'Linn. Phil. Bot.,' Sec. 124. + +[115] 'Bot. Zeit.,' 1857, p. 873. See also 'Verhandl. Nat. Hist. +Vereins. Preuss. Rheinl. u. Westphal.,' 1854, t. ix. + +[116] "Pannicula spicatim sparsa onusta innumera foetura herbaceorum +flosculorum racematim cohaerentium," 'Lobel. Stirp. Hist.,' p. 163. This +is the "Besome Plantain, or Plantain with spoky tufts," of Ray, +'Synopsis,' p. 314. Gerard's 'Herbal,' Ed. Johnson, p. 420. Parkinson, +'Theat. Bot.,' p. 494. Baxter, 'Loudon. Mag. Nat. Hist.,' vol. ix. p. +204, and vol. iii, p. 482. fig. 118. + +[117] 'Flora.' 1856. p. 706. + +[118] 'Flora of Berwick-on-Tweed,' vol. i. p. 38. + +[119] 'Adansonia,' vol. iv. 1864, p. 150, tab. vii. 'Gard. Chron.,' +November 19th, 1864. + +[120] 'Ann. Sc. Nat.,' ser. 3, tom. ii, p. 290; and 'Adansonia,' iii, +tab. iv; see also Bureau, in 'Bull. Soc. Bot. France,' x, p. 191. + +[121] Baillon, 'Adansonia,' i, 286. + +[122] See also figure in 'Hort. Eystett. Ic. Plant. Vern.,' fol. 15, +fig. 1. _Ranunculus asiaticus_. + +[123] Duchartre, 'Ann. des sc. nat.,' 3me serie, vol. ii, 1844, p. 293. + +[124] Reissek, 'Linnaea,' vol. xvii, 1843, p. 641, tab. xix. + +[125] The tube of the calyx in these specimens was traversed by ten +ribs, apparently corresponding to the primary ridges of the normal +fruit; these ribs were destitute of spines, and the bristly secondary +ridges were entirely absent. Those portions of the carpels which were +detached from the calyx had each three ribs, a central and two lateral +ones, which appeared to be continuous with the ribs of the calyx +below,--although in the case of the calyx there were ten, in the case of +the carpels six ribs, three to each. This diversity in number is thus +explained:--A circle of vascular tissue ran round the interior of the +calyx-tube, at its junction with the limb, and at the point of insertion +of the petals and stamens. The vascular circle seemed to be formed from +the confluence of the ten ribs from below. Of the five ribs in each half +of the calyx, the three central ones were joined together just at the +point of confluence with the vascular circle, above which they formed +but a single rib--that traversing the centre of the carpellary leaf; the +two lateral ribs of each half of the calyx seemed to be continuous, +above the vascular rim, with the lateral ribs of the carpel; these +lateral ribs were connected on either side with the central one by short +branches of communication. The disposition of the ten ribs may be thus +represented:-- + + 1 1 1 1 1 1 + 3 2 3 2 3 3 2 3 2 3 + 1 1 1 1 1 1 1 1 1 1 + +The lower line of figures represents the calycine ribs, the middle row +shows how each of these ribs is divided at the vascular rim, and the +uppermost row shows their distribution above the rim. From this it will +be seen that six of the calycine ribs divide into three branches, one +prolonged upwards as a lateral or median rib into the carpellary leaf, +the other running horizontally to join with similar branches sent out +from the neighbouring rib; the four intermediate calycine ribs divide +into two branches only, which join the side branches of the first +mentioned, but have no direct upward prolongation into the carpel. The +ten ridges are placed opposite to the sepals and petals. + +[126] 'Neue Denkschriften der allgemeine Schweizerischen Gesellschaft,' +band 5. 1841. tab. 2. + +[127] Bell Salter, 'Gard. Chron.,' March 13th, 1847, and 'Ann. Nat. +Hist.,' 1847, vol. xix, p. 471. &c. + +[128] 'The Origin and Production of Proliferous Flowers, with the +Culture at large for raising Double Flowers from Single, and Proliferous +from the Double.' By J. Hill, M.D. London, 1759. + +[129] A. de Candolle, 'Neue Denkschriften,' op. cit., p. 9; also Unger +as cited in 'Botanical Gazette,' May, 1351. p. 70. + +[130] Duchartre, op. cit. + +[131] 'Ann. Sc. Nat.,' 1844, vol. i, p. 297. + +[132] Maout, 'Lecons Elementaires de Botanique,' vol. ii. p. 488; +Ferrari. 'Hesperides.' pls. 271, 315, 405. + +[133] Moquin-Tandon, loc. cit., p. 386, &c.; see also Trecul, in the +'Bull. Soc. Bot. France,' tom. i, p, 307. + +[134] 'Bull. Soc. Bot. Fr.,' 1863, vol. x, p. 461. + +[135] 'Ann. Sc. Nat.,' 1835, p. 65. See also Le Maout, 'Lecons +Element.,' vol. ii, p. 426. + +[136] 'Proc. Hort. Soc.,' vol. i, p. 39, fig. 2. + +[137] See also 'Nat. Hist. Review,' 1865, p. 377. + +[138] 'Acad. Roy. Belg.,' April 11th. 1863. + +[139] 'Bull. Soc. Bot. Fr.,' tom. iii, 1856, p. 479. + +[140] 'Linnaea,' vol. xv, p. 266, _c. ic._ Caspary, 'Schriften d. +Physik.-Oek. Gesell. zu Koenigsberg,' bd. ii, p. 5, tab. iii, fig. 39, +&c. + +[141] Lindley, 'Veg. King.,' p. 545; also Clarke on the Position of +Carpels, Linn. Soc.,' December, 1850. 'Proc. Linn. Soc.,' ii, p. 105. + +[142] 'Notulae,' vol. i, Dicot. p. 127. 'Atlas,' pl. xliii. + +[143] Moquin-Tandon gives the following references to cases of +proliferous roses, but some I have not been able to verify. 'Journ. des +Sav.,' 22 Mai 1679. Hottinger, 'Ephem. Nat. Cur.,' dec. 3 ann. 9 et 10, +p. 249. Marchant, 'Mem. Acad. Scienc. Paris.' 1707, p. 488. Preussius, +'Ephem. Nat. Cur.,' cent. 7 et 8. App. p. 83. Schuster, 'Act. Acad. Nat. +Cur.,' vol. vi, p. 185. Spadoni, 'Mem. Soc. Ital.,' t. v, p. 488. See +also at the end of this section for numerous other references. + +[144] 'Bull. Acad. Belg.,' t. xx, part ii, p. 271. See also Bellynck, +'Bull. Soc. Bot. Belg.,' t. vi, ex. 'Bull. Soc. Bot. France,' t. xiv, +1867, Rev. Bibl., p. 241. _Orchis ustulata_. + +[145] I have not been able to meet with this, but it is said to contain +a paper on prolification, with numerous bibliographical references. + +[146] 'El. Ter. Veg.,' p. 364, Adnot. + + + + +CHAPTER III. + +HETEROTAXY. + + +Under this category are here included a variety of deviations from the +ordinary arrangement and position of parts which cannot conveniently be +classed under the preceding or under other headings. The term heterotaxy +is intended to apply to the production of organs in situations where, +under usual circumstances, they would not be formed. It thus does not +include cases of substitution, where one part is replaced by another, or +more or less metamorphosed, nor cases of multiplication, nor of +prolification which are characterised not only by the production of +members in unwonted situations, but also in unwonted numbers. From the +very nature of the anomalies, and specially from the scanty knowledge we +possess concerning their mode of development, it is not possible to +allocate them in all cases correctly, and moreover many of them might as +well be placed in one group as in another. + +=Formation of adventitious roots.=--This is of exceedingly common +occurrence in a vast number of plants, so much so that in most cases it +cannot be considered as in any way abnormal; there are, however, a few +instances where the formation of these organs may be considered to come +within the scope of teratology, or, at least, where their production is +the result of injury or of some unfavorable condition to which the plant +is exposed. + +Thus the production of adventitious roots on the stem of the vine is +considered to be due to untoward circumstances impairing the proper +action of the ordinary subterranean roots. So, too, the formation of +roots on the upper portions of stems that are more or less decayed +below, as in old willows, is to be considered as an attempt to obtain +fresh supplies through a more vigorous and healthy channel. + +A similar occurrence often arises as a consequence of some injury. +Virgil had this circumstance in view when he wrote + + "_Quin et, caudicibus sectis, mirabile dictu, + Truditur e sicco radix oleagina ligno._"--'Georg.' Bk. ii. + +I have seen many specimens of adventitious roots produced on the olive +in the way just mentioned. + +In the 'Gardeners' Chronicle,' January 8th, 1853, p. 21, is described a +curious formation of roots in the fissure between two divisions of a +laburnum stem. In the same journal, January 1st, 1853, p. 4, Mr. Booth +mentions the case of a Cornish elm, the trunk of which was divided at +the top into two main divisions, and from the force of the wind or from +some other cause the stem was split down for several feet below the +fork. Around the edges of the fracture, layers of new bark were formed, +from which numerous roots issued, some measuring an inch in diameter and +descending into the cleft portion of the tree: similar instances must be +familiar to all observers. + +It may happen that these roots sent down into the cavity of a decaying +trunk may, after a time, become completely concealed within it, by the +gradual formation and extension of new wood over the orifice of the +cavity formed by the death and decay of the old wood. Such is presumed +to be the explanation of a specimen of this kind in the possession of +the writer, and taken from a cavity in an apparently solid block of +rosewood; externally there were no marks to indicate the existence of a +central space, but when the block was sawn up for the use of the +cabinet-maker, this root-like structure was found in the centre and +attached to one end of the cavity. + +The production of roots which ultimately serve as props to support the +branches, or as buttresses to compensate for the increasing weight of +branches and foliage, is also a familiar occurrence. The huge gnaurs and +burrs met with occasionally on some trees often produce great +quantities, not only of adventitious buds, but of roots also. + +[Illustration: FIG. 71.--Production of adventitious roots from leaf +stalk of celery.] + +The leaves, equally with the stems, have the power of emitting roots +under certain conditions, as when the leaves are in close contact with +moist soil or as the result of injury. This happens in some plants more +readily than in others--_Bryophyllum calycinum_ is a well-known +instance. Mr. Berkeley has described the formation of roots from the +fractured leaves of celery,[147] and also in a cabbage where a snail +"having gnawed a hole into the middle of a leaf at its junction with the +stem, a fascicle of roots was formed, bursting through the tissue lining +the cavity, and covered with abundant delicate hairs after the fashion +of ordinary radicles." + +[Illustration: FIG. 72.--Germinating plant of mango, showing production +of roots from one of the cotyledons (from the Kew Museum).] + +The production of adventitious roots is not limited to the ordinary +leaves of the plant, but may be manifested on the cotyledons; thus +Irmisch describes cases of this kind in the cotyledons of _Bunium +creticum_ and _Carum Bulbocastanum_.[148] I have figured and described +an analogous case in the cotyledons of the Mango (fig. 72).[149] + +To this formation of adventitious roots the gardener owes the power he +has of propagating plants by cuttings, _i.e._, small portions of the +stem with a bud or buds attached, or in some cases from portions of the +leaves, of the roots themselves, or even of the fruit, as in the case of +the cactus (Baillon). Care also has to be exercised in grafting certain +fruit trees not to allow the grafted portion to be too close to the +ground, else the scion throws out roots into the soil, and the object of +the cultivator is defeated. + +[Illustration: FIGS. 73 and 74 show formation of roots from leaves +induced by the art of the gardener.] + +Layering is another garden operation dependent on the formation of these +organs, and advantage is also sometimes taken of this tendency of some +plants to produce roots when injured to reduce the dimensions of a plant +when getting too large for the house in which it is growing. By +gradually inducing the production of new roots from the central or upper +portions of the stem, it becomes possible, after a time, to sever the +connection between the original roots and the upper portion of the +trunk, and thus secure a shortened plant. + + On the subject of adventitious roots, &c., reference may be + made to Trecul, 'Ann. Sc. Nat.,' 1846, t. v, p. 340, et vi, p. + 303. Duchartre, 'Elements de Botanique,' p. 219. Lindley, + 'Theory and Practice of Horticulture.' Thomson's 'Gardener's + Assistant,' pp. 374, _et seq._; and any of the ordinary + botanical text-books. + +=Formation of adventitious buds on roots.=--One of the characteristics +by which roots are distinguished from stems in a general way consists in +the absence of buds; but, as is well known, they may be formed on the +roots under certain circumstances, and in certain plants, e.g., _Pyrus +Japonica_, _Anemone Japonica_, &c. What are termed suckers, owe their +origin to buds formed in this situation. + +If roots be exposed or injured, they will frequently emit buds. The +well-known experiment of Duhamel, in which a willow was placed with the +branches in the soil and the roots in the air, and emitted new buds from +the latter and new roots from the former, depended on this production of +adventitious organs of either kind. + +Gardeners often avail themselves of the power that the roots have of +producing buds to propagate plants by cuttings of the roots, but in many +of these cases the organ "parted" or cut is really an underground stem +and not a true root. + +M. Claas Mulder has figured and described a case in the turnip-radish of +the unusual formation of a leafy shoot from the root, apparently after +injury.[150] From the figure it appears as if the lower portion of the +root had been split almost to the extremity, while the upper portion +seems to have a central cavity passing through it. From the angle, +formed by the split segments below, proceeds a tuft of leaves, some of +which appear to have traversed the central cavity and to have emerged +from the summit, mingling with the other leaves in that situation. The +production of a flower-bud has even been noticed on the root of a +species of _Impatiens_. + +=Formation of shoots beneath the cotyledons.=--The tigellar or axial +portion of the embryo plant, as contrasted with the radicle proper, is +very variously developed in different cases; sometimes it is a mere +"collar" bearing the cotyledons, while at other times it is of +considerable size. Generally it does not give origin to shoots or leaves +other than the seed-leaves, but occasionally shoots may be seen +projecting from it below the level of the cotyledons. This happens +frequently in seedling plants of _Anagallis arvensis_, _Euphorbia_ +_peplus_, and other species, _Linaria vulgaris_, some _Umbelliferae_, +&c.[151] + +=Adventitious formation of leaves.=--The term phyllomania has been +vaguely applied both to the production of an unwonted number of leaves +and to their development in unusual situations. Under the present +heading the latter class of cases are alone included. The extraordinary +tendency in some Begonias to develop leaves or leafy excrescences from +their surfaces is elsewhere alluded to, and is, in reality, a species of +hypertrophy or over-luxuriant growth. + +In some flowers where the inferior ovary is supposed to be, in part at +least, formed by a dilatation of the top of the flower-stalk, leaves +have been met with proceeding from the surface of the ovary or fruit, as +in _Crataegus tanacetifolia_, roses, pears, gooseberries, &c. In a +specimen of _Nymphaea alba_ I have met with scale-like leaves projecting +from the surface of the fruit (or torus?), and which did not appear to +be metamorphosed stamens or styles (fig. 76). + +[Illustration: FIG. 75.--Leaf proceeding from hip of the Rose.] + +[Illustration: FIG. 76.--Leaves proceeding from the ovary of +_Nymphaea_.] + +For other illustrations of increased leaf-formation, see Multiplication +of foliar organs. + +[Illustration: FIG. 77.--_Leontodon_. Scape with two leaves; the bracts +of the involucre are also leafy.] + +=Production of leaves on a usually leafless inflorescence.=--The +development of the bracts of an inflorescence to such an extent that +they resemble ordinary leaves is elsewhere alluded to as of common +occurrence. It happens far less frequently that leaves are developed on +an inflorescence usually destitute of them, without any metamorphosis or +substitution, and without any formation of adventitious buds, such as +happens in prolification. Such a partial change from a floriferous to a +foliiferous branch may be seen in a specimen of _Sambucus nigra_ in the +Smithian herbarium in the Linnean Society, where the ultimate branches +of the cyme bear small leaves. My attention was directed to this +specimen by the Rev. W. Newbould. + +Jacquin figures an analogous case in _Sempervivum sediforme_,[152] in +which the branches of the inflorescence were prolonged into leafy +shoots. + +Sometimes from the side of a flower-stalk or scape, which usually does +not bear leaves, those organs are produced. The common dandelion, +_Taraxacum_, sometimes offers an illustration of this, and also the +daisy (_Bellis_).[153] In a specimen of fasciated cowslip given me by +Mr. Edgeworth there was a similar formation of leaves on the flattened +stalk. + +=Production of leaves or scales in place of flower-buds.=--The position +of the leaf and of the flower-buds respectively is, in most plants, well +defined, but occasionally it happens that the former is formed where, +under ordinary circumstances, the latter organ should be. This may +happen without the formation of any transitional organs between the two, +and without actual increase in the number of the buds. Where there is +evidently a passage from leaf-bud to flower-bud, or _vice versa_, the +case would be one of metamorphy. If the number of buds be augmented, or +they be mixed with the flower-buds, then it would be referable to leafy +prolification of the inflorescence. There remains a class of cases +wherein there is a complete substitution of one structure for the other, +it may be without the slightest indication of transition between the +two, and without any admixture of leaf-buds among flower-buds, or any +absolute increase in the number of organs, as in Prolification. Such a +case is represented in fig. 78, which shows a portion of the stem of a +species of _Valeriana_, bearing at the summit, not an inflorescence, but +a tuft of leaves without the slightest indication of flowers. + +Drs. Hooker and Thomson relate that in Northern India the flowers of +_Anemone rivularis_ are very generally absent, and their place supplied +by tufts or umbels of leaves.[154] In the collection of the late Mr. N. +B. Ward was a specimen of lupin in which the flowers were all absent, +and their place supplied by tufts of leaves. + +[Illustration: FIG. 78.--Tuft of leaves replacing the inflorescence in a +species of _Valeriana_.] + +A similar appearance has been noticed in _Compositae_, and I owe to the +kindness of Professor Oliver the communication of a specimen of a +species of _Bidens_ from Peru, in which the capitula, instead of +consisting of florets, as usual, contained tufts of linear ciliolated +bracts within the involucre, without a trace of flowers. In the eleventh +volume of the 'Linnaea,' 1837, p. 301, Von Cesati figures and describes +an analogous case in _Carduus crispus_. The same author[1] records a +similar instance in the umbel of _Seseli coloratum_, where the place of +the flowers was occupied by stalked tufts of leaves. In the 'Gardeners' +Chronicle,' October 6th, 1860, p. 894, is mentioned an instance where +the blossoms of the pea were entirely absent, and their place supplied +by accumulations of small, ovate, green scales, thus presenting an +appearance similar to that brought about by the inordinate +multiplication of the sepals in the "wheat-ear carnation," and in the +Sweet William, and not unlike the condition met with in _Bryophyllum +proliferum_. In _Digitalis purpurea_ a similar anomaly is sometimes met +with. + +In the apple I have observed leafy shoots bearing terminal tufts of +leaves where the flower should have been, so that what, under ordinary +circumstances would be a corymb of flowers, is here represented by a +series of tufts of leaves. In the cultivated azaleas also, leafy shoots +occupying the position of the flower may occasionally be met with. + +In _Bouchea hyderabadensis_ I have seen the inflorescence more than +usually branched and covered with little tufts of bracts, without a +trace of true flower. A similar condition seems not infrequent in +_Gentiana Amarella_, as I have not only met with the plant myself in +this condition, but have been favoured with specimens by Mr. Pamplin, +Mr. Darwin, and others. In _Phyteuma spicatum_ an analogous appearance +has been recorded. + +Among Griffith's collections from Affghanistan is a species of willow +(_Salix_) in which the inflorescence replaced by a much branched +panicle, bearing a quantity of minute bracts, in the axils of which +nestle numerous small buds. In another specimen the inflorescence +preserves its usual catkin-like shape, but the flowers are replaced by +little tufts of leaves. M. Germain de Saint Pierre mentions a case +wherein the flowers of _Alisma parnassifolia_ were completely replaced +by leaf-buds.[155] + +[Illustration: FIG. 79.--Spikelets of _Willedenovia_, composed entirely +of scales to the exclusion of flowers.] + +[Illustration: FIG. 80.--Rose Willow, _Salix_, sp.] + +Here, also, may be mentioned the curious aggregations of scales which +occur in some grasses, in _Restiaceae_, _Juncaceae_, and other orders, in +which the inflorescence is made up of collections of scales or bracts +with no trace of floral structure. Fig. 79 shows this in a species of +_Willdenovia_, and a very good example is figured in a bamboo, +_Pseudostachyum polymorphum_, by General Munro.[156] + +"Rose willows" (fig. 80) owe their peculiar appearance to a similar +cause, the scales of the catkin being here replaced by closely crowded +leaves. These aggregations of scales or leaves are not confined to the +inflorescence, but may be found in other parts of the plant, and may be +frequently met with in the willow, birch, oak, &c., generally as the +result of insect puncture. On the other hand, the production of leaves +or leaf-buds in place of flowers is, as is well known, generally the +consequence of an excess of nutrition, and of the continuance rather +than of the arrest of vegetative development.[157] It has even been +asserted that a flower-bud may be transformed into a leaf-bud by +removing the pistil at a very early stage of development, but this +statement requires further confirmation.[158] + +=Viviparous plants.=--The spikelets of certain grasses are frequently +found with some of their constituent parts completely replaced by +leaves, like those of the stem, while the true flowers are usually +entirely absent. A shoot, in fact, is formed in place of a series of +flowers. In these cases it generally happens that the outermost glumes +are changed, sometimes, however, even the outer and inner paleae are +wholly unchanged, while there is no trace of squamulae or of stamens and +pistils within them, but in their place is a small shoot with miniature +leaves arranged in the ordinary manner. + +The grasses most commonly affected in this manner are _Dactylis +glomerata!_, _Poa bulbosa!_, _Poa annua!_, _P. trivialis!_, +_pratensis!_, _alpina!_, _angustifolia_, and _laxa_, _Cynosurus +cristatus_, _Festuca nemoralis_, _F. ovina!_, _Glyceria fluitans!_, _Gl. +aquatica_, _Aira alpina!_, _caespitosa!_, _Phleum phalaroides_, _Lolium +perenne!_, _Alopecurus pratensis!_, _Agrostis alba_, _Holcus mollis!_ + +[Illustration: FIG. 81.--Portion of panicle of _Aira vivipara_ and +separate floret.] + +From an examination of the structure of viviparous grasses Von Mohl was +led to the conclusion that the lower palea is to be considered as a +bract, and not a perianthial leaf, because the base of the palea +surrounds the stem or axis of the spikelet entirely, and both its +margins cohere towards its lower extremity.[159] + +A similar condition occurs not infrequently in _Polygonum viviparum_, +and in _Juncaceae_, _Cyperaceae_, &c. + +In the genus _Allium_ an analogous formation of little buds or bulbils +takes place in lieu of flowers; this is specially the case with _A. +vineale_, the flowers of which are rarely seen. + +Other illustrations of a similar character, where the adventitious +leaf-buds are mixed in amongst the flower-buds, are cited under the head +of Prolification of the Inflorescence. + +=Formation of buds on leaves.=--The formation of little bulbs upon the +surfaces or edges of leaves, forming what are called viviparous leaves, +has long been familiar to botanists amongst Alliums. Professor Alexander +Braun,[160] who has paid much attention to this subject, divides cases +of this kind according to the position of the buds; thus, for instance, +they are sometimes formed upon the upper portion of the leaf or petiole, +as in many ferns, in _Nymphaea guineensis_, some _Arads_, &c. The same +condition has been met with as a teratological occurrence in the leaves +of _Cardamine pratensis_, _Hyacinthus Pouzolzii_, _Drosera +intermedia_,[161] _Arabis pumila_, _Chelidonium majus_, _Chirita +sinensis_,[162] _Episcia bicolor_,[163] _Zamia_, &c.[164] Many species +of _Begonia_ possess the power of emitting buds from the petioles and +veins of the leaf; the little ramenta or scales which so plentifully +beset the surface of some of these plants likewise, in some instances, +pass gradually into leaves. _B. phyllomaniaca_, Mart., is the species +best known as manifesting this tendency, but others have it also.[165] + +Buds are also very often formed upon the margins of the leaf, the best +known instance of which occurs in _Bryophyllum calycinum_; +Weinmann[166] figures an instance of this kind in _Alchemilla minima_, +or they may occur upon the lower surface of the leaf, as in +_Ornithogalum scilloides_ and _longe-bracteatum_. M. Duchartre[167] +mentions a case in the tomato in which the leaves gave origin to small +leaf-bearing branches, which, of course, must have originated from buds, +just in the same way as in the _Drosera_ before mentioned. + +[Illustration: FIG. 82.--Formation of shoot on leaf of _Episcia +bicolor_.] + +Gardeners occasionally avail themselves of this formation of buds from +leaves to propagate plants, _e.g._ _Hoya_, _Gesnera_, _Gloxinia_, &c. + +=Formation of buds in the pith.=--This is said to be a normal condition +in the curious _Stangeria paradoxa_,[168] and Mr. Berkeley records an +instance of this in sea-kale[169] (fig. 83) where the crown had been +injured, and buds were seen sprouting from its centre. + +[Illustration: FIG. 83.--Adventitious buds in sea kale.] + +[Illustration: FIG. 84.--Hyacinth bulb cut across to induce the +formation of new bulbs.] + +[Illustration: FIG. 85.--Showing the formation of new bulbs on the cut +edges of an old hyacinth bulb.] + +It will be remarked that the adventitious production of buds, like that +of roots, is very often consequent on decay or injury. The Dutch +bulb-growers have availed themselves of this latter circumstance in the +propagation of hyacinths. Mr. Fortune, who published some articles on +this subject in the 'Gardener's Chronicle,'[170] describes two special +modes as adopted by these skilful horticulturists--the one to make two +or three deep cuts at the base of the bulb, destroying the nascent +flower-stalk when, after a time, small bulbs are formed along the edges +of the cut surfaces (figs. 84, 85). The other method is effected by +scooping out the interior of the base of the bulb, thus leaving exposed +the cut ends of the sheathing leaves arranged concentrically; along +these lines the new bulbs are, after some time, formed in great numbers +(fig. 86). + +[Illustration: FIG. 86.--Showing the production of small bulbs on the +inner surface of the scooped-out bulb of hyacinth.] + +For the formation of supernumerary leaves on the surface of the normal +one, see Multiplication and Hypertrophy. + +=Production of gemmae in place of spores.=--An instance of this is +recorded by Dr. Montagne[171] in the case of a moss, _Encamptodon +perichaetialis_, in which, in the interior of the capsule, in lieu of +spores numerous minute gemmae of the same nature as those in the cup of +_Marchantia_ were seen. + +=Formation of flowers on leaves.=--It is very doubtful whether a +flower-bud has ever been found actually on a leaf. Mere adhesion of the +pedicels of the leaf, such as happens in _Ruscus_, in _Helwingia_, +_Erythrochiton hypophyllanthus_, and a few other plants, is, of course, +not really to be considered in the light of an actual growth from the +leaf, and it is very doubtful in the present state of our knowledge +whether the case of the Nepaul barley should find a place here, but for +convenience sake it is placed in this section, as it is uncertain at +present where it properly belongs. + +[Illustration: FIG. 87.--Three-lobed end of outer palea of Nepaul barley +bearing supplementary florets.] + +[Illustration: FIG. 88.--Three spikelets of Nepaul barley.] + +[Illustration: FIG. 89.--Lip of outer palea of Nepaul barley.] + +[Illustration: FIG. 90.--Supplementary rachillus or outer palea of +Nepaul barley bearing florets.] + +[Illustration: FIG. 91.--Diagram showing arrangement of supplementary +rachillus and florets.] + +[Illustration: FIG. 92.--Supplementary floret of Nepaul barley; palea +removed.] + +This curious plant has been described and figured by Irmisch in the +13th volume of the 'Linnaea,' p. 124, t. iv; also by Professor Henslow, +'Hooker's Journal of Botany,' 1849, vol. i, p. 33, tabs. 2, 3. The lower +palea of this plant forms an inverted flower-bud upon its midrib. In +some fresh specimens which I have lately examined I find the structure +to be as follows:--On each notch of the rachis there are three spikelets +(fig. 88), each one-flowered, and each provided with two linear glumes; +the outer palea in all cases is three-lobed at the summit, the central +lobe being oblong and hollow, forming a kind of hood (figs. 87-89), and +covered with hairs, which are directed downwards towards the centre of +the plant. The two lateral lobes are more pointed than the central one; +like it they are provided with hairs, but the hairs, in this case, are +turned away from the centre of the plant. The cavity of the side lobes +is generally empty, but that of the central lobe is occupied by a very +slender stalk, which is apparently the termination of the midrib, but +which is bent inwards at an acute angle, so as to occupy the hollow +space (figs. 90-91). On this slender axis are developed two florets, +more or less imperfect in their structure. Only one of the florets that +I have seen contained a perfect ovary. The tips of the lateral lobes of +the paleae in the primary flower are sometimes extended into a long awn. +A similar awn may also be occasionally found on the tips of the paleae of +the rudimentary florets. The occurrence of an adventitious axial +structure with rudimentary flowers has been adduced in support of the +opinion that the lower paleae is, at least so far as its midrib is +concerned, an axial rather than a foliar structure, but in the present +uncertain state of our knowledge as to the morphology of grasses it is +hazardous to risk any explanation founded on so exceptional a case as +that of the Nepaul barley.[172] + +=Production of flower-buds in place of leaf-buds.=--Under natural +circumstances this does not appear to be of so common occurrence as the +change above alluded to, but by the art of the gardener the change is +often effected. In rhododendrons and in peach trees and roses I have met +with this change occurring without human agency. The means adopted by +the gardener are such as check the luxuriance of the leaf-shoots,[173] +and this is effected in various ways, as by continuous "pinching" or +removal of the leaf-buds, by pruning, ringing the bark, confining the +roots, limiting the supply of nutriment, and other means all based on +the same principle. Some of the Cape bulbs (_Cyrtanthus_) are known not +to produce their flowers till their leaves have received, in some +manner, a check. Fires which often destroy the herbage thus have the +effect of throwing the plant into bloom. A very remarkable instance is +recorded of the production of flower-buds after an injury to the +leaf-buds in the 'Bulletin of the Botanical Society of France,' vol. ix, +p. 146. It appears that during the war of the French against the Arabs +in Algiers, the latter planted several hundreds of Agaves with a view to +obstruct the passage of the French cavalry. The soldiers hacked these +plants with their sabres, and cut out the central tuft of leaves, or the +heart, as gardeners call it. The following season almost every one of +these Agaves sent up their large handsome flower-spikes. It is well +known that, under ordinary circumstances, these plants do not flower +except at long intervals of time. + +=Presence of flowers on spines.=--That the spine, as a contracted +branch, should occasionally produce flowers is not to be wondered at, +though the occurrence is by no means common. M. Baillon showed at a +meeting of the Botanical Society of France ('Bulletin,' vol. v, 1858, p. +316) a branched spine of _Gleditschia_ bearing a flower at the end of +each of the sub divisions. This was, therefore, strictly analogous with +those cases in which the peduncle is normally spiney. + +=Formation of flower-bud on the petals.=--An instance of this, it is +believed, the only one on record, is cited in the 'Gardeners' Chronicle' +for 1865, p. 760, by the Rev. M. J. Berkeley, who describes the +formation of a flower-bud on the surface of a petal of _Clarkia +elegans_. Reasoning from analogy there seems no reason why buds should +not be formed on the petals as well as on the leaves. + +=Formation of buds on fruits.=--This is a point of some moment with +reference to the share which the axis takes in the production of +"inferior" fruits. A very frequent malformation in pears is one wherein +a second pear proceeds from the centre of the first, and even a third +from the centre of the second.[174] Pears are occasionally also observed +arising either from the axils of the sepals of the primary pear or from +the axil of leaves originating on the outer surface of the fruits--using +the term fruit in its popular sense. These cases afford strong +confirmation of the view that the outer portion of the so-called fruit +in these plants is rather to be considered as an expansion and +hollowing-out of the flower-stalk, than as formed from the calyx-tube. +It is noteworthy that the true carpels and seeds are frequently entirely +absent in these cases.[175] Further reference to these fruits will be +made under the head of Hypertrophy. + +M. Trecul has described and figured an instance in a species of +_Prismatocarpus_, in which a second flower proceeded from the axil of a +bract attached to the side of the fruit of the first flower.[176] A +similar growth was observed in the fruit of _Philadelphus speciosus_ by +M. A. Gris, who observed that the so-called calyx-tube was provided with +two small bracts, from the axil of one of which proceeded a small +flower-bud.[177] + +[Illustration: FIG. 93.--Small buds projecting from the edges of the +fruit in _Opuntia_.] + +The fruits of _Opuntia Salmiana_, _O. fragilis_,[178] _O. monacantha_, +and of some species of _Echinocactus_, have been observed to form small +fruit-like branches around their summits. M. Napoleon Doumet describes +the fruit as ripening as usual, but as being destitute of seeds in the +interior; after a little while the fruit begins to wither, and then a +circle of small buds, like those of the stem, may be seen at the top of +the fruit, each bud springing from the axil of a little tuft of wool and +spines found on the fruit. These little buds elongate into long shoots, +produce flowers the following year, which flowers exhibit the same +peculiarity. Gasparini and Tenore are said to have recorded the same +fact as long since as 1832. The specimen from which the figure (fig. 93) +was taken produced its fruits in the Royal Gardens at Kew, and is now +preserved in the museum of that establishment. The adventitious growth +in these cases appears to arise from the tufts of spines, which, it has +been suggested, are the homologues of the sepals. There can, however, be +little doubt that the outer and lower portion of the fruit of _Opuntia_ +and its allies is a dilatation of the flower-stalk. This is borne out by +the fruits of _Pereskia_, which bear leaves on their surface arranged +spirally; indeed, the fruits of _Pereskia Bleo_ are mentioned as +producing buds from their summits, in the same way as the _Opuntia_ +just cited. _P. Bleo_ is said, by M. Delavaud,[179] to present this +anomaly as a constant occurrence. On the summit of the primary fruit, +arising apparently from the axils of the sepals, or of small leafy +bracts in that situation, are a series of fruit-like branches, which, in +their turn, are surmounted by others, even to the fourth generation. + +The fruits of _Tetragonia expansa_ frequently have attached to their +side a secondary flower or fruit in such a position as to lead to the +inference that it springs from the upper portion of the peduncle which +is dilated to invest the true carpels. In other instances it is due to +an adhesion of the pedicel to the side of the fruit. In either case the +production of an adventitious bud might be considered as an illustration +of prolification of the inflorescence, though not as was supposed by +Moquin and others of axillary prolification.[180] + +Buds have also been produced artificially on the surface of some of the +fruits in the construction of which the axis is supposed to share; thus, +the unripe fruits of some species of _Lecythis_ were stated by Von +Martius, at a meeting of the German Naturalists at Carlsruhe, to produce +buds when placed in the earth. The fruit of these plants is probably of +the same nature as that of the _Pomaceae_, and Baillon[181] succeeded in +producing buds on the surface of the inferior ovary of _Jussiaea_. + +Some of the cases just mentioned have been considered to be instances of +prolification of the fruit, but the fruit has little to do with the +appearances in question. + +=Formation of adventitious flowers and fruits within the ovary.=--This +generally arises either from substitution of a flower-bud for an ovule +or from prolification; there are certain cases, however, where the new +growth seems not to be either due to metamorphosis or to prolification +strictly. + +The cut, fig. 94, represents a case where, in the dilated upper portion +of the ovary of _Sinapis arvensis_, two flower-buds were found +projecting from a raised central line, corresponding, as it would seem, +to the midrib, and not to the margins of the carpel. Similar cases have +occurred in _Nasturtium amphibium_, _Brassica Rapa_, and _Passiflora +quadrangularis_. + +[Illustration: FIG. 94.--Distended pod of _Sinapis arvensis_ bearing in +the interior stalked flower buds.] + +In Bromfield's 'Flora Vectensis,' p. 35, the following account is given +of an abnormal development in _Cardamine pratensis_: "On the lower part +of the corymb were several seed vessels on pedicels changed from their +usual linear to an ovate elliptical figure, so as to resemble a +silicula. These, on being opened, were found to contain petals of the +usual colour, which in the pods above had burst from their confinement +and appeared as semi-double flowers; the valves of the pod answering to +the true calyx. * * * From their verticillate arrangement it is evident +that these petaloid expansions were not transformed seeds, but simply a +development of the common axis within the ovary into an abortive whorl +of floral organs, besides which there were evident rudiments both of +stamens and germens in the centre of the bundle." Baillon[182] also +records a case of the same nature in _Sinapis arvensis_. + +[Illustration: FIG. 94*.--Portion of the interior of the silicle in +_Cheiranthus Cheiri_, showing adventitious pod in the place of an +ovule.] + +[Illustration: FIG. 95.--Adventitious pod from fig. 94, enlarged.] + +Here, too, may also be mentioned the presence of an adventitious siliqua +within the ordinary one attached along the same line as the ovules, and +partially divided by a replum into two cavities. In this case there was +nothing to indicate the presence of floral envelopes (figs. 94, 95). A +similar occurrence has been brought under my notice in some grapes which +were observed to be cracking before they were perfectly ripe, and in +which adventitious fruits were found within the parent grape, occupying +the position of seeds (figs. 96, 97). + +Similar anomalous growths are noticed under the heads of Substitution +and Prolification. + +=Formation of stamens within the cavity of the ovary.=--The only +instance of this that has come under the author's observation occurred +in some flowers of _Baeckea diosmaefolia_, Rudge, for an examination of +which he is indebted to Mr. Bentham. + +[Illustration: FIG. 96.--Section of Barbarossa grape showing +adventitious grape in the position of a seed.] + +[Illustration: FIG. 97.--Grape with supplementary fruit in the interior] + +In the normal flower there is a turbinate hollow calyx, whose limb is +divided into five serrated lobes; alternating with these latter, and +springing from the throat of the calyx, are the petals. Originating from +the same annular disk as the petals are the stamens, seven or eight in +number. The ovary is partially adherent, is surmounted by a style, and +has two or three loculi with an axile placenta, to which several small +curved ovules are attached. The malformed flowers did not present +anything peculiar in their outer parts, nor did the ovary, partially +immersed within the expanded top of the flower-stalk and the calyx-tube, +which is continuous with that organ, show externally any indication of +the change within. On cutting it across, however, in any direction, +numerous perfect stamens (filaments and anthers) were seen projecting +from the walls of the cavity (fig. 98). In most of the flowers the ovary +was one-celled; but in a few there was the usual axile placenta; yet +even in these latter cases the stamens originated from the walls of the +cavity, and not from the placenta. The stamens presented different +degrees of development; in some cases they were fully formed, the +anther-lobes open, and the pollen exposed; while in other instances the +filaments were involute or circinate, just as the ordinary stamens are +in the unexpanded flower-bud. In some cases imperfect stamens were +found, mere barren filaments, with or without rudimentary anthers at the +top. In no instance was there a perfect ovule, or, indeed, any trace of +ovules. The stamens appeared to be arranged irregularly on the walls of +the ovarian cavity; and while they were certainly more numerous at the +lower portion (that now generally considered to be formed by the +cup-like end of the pedicel), they were not wanting in the upper half of +the ovary (or that which is probably formed from the carpellary leaves). + +[Illustration: FIG. 98.--1. Vertical section of flower of _Baeckea +diosmaefolia_, showing stamens within the ovary; magnified ten times. 2. +Transverse section of ovary. 3. Stamen. 4. Imperfect stamen.] + +This case differs from most that have been recorded, and in which there +has been a more or less complete substitution of anther for carpel, or +where the tissues of the carpel have produced pollen, and so taken upon +themselves the appearance and functions of anthers. Instances of this +latter kind are not uncommon; but in the _Baeckea_ there were perfect +stamens proceeding from perfect and completely closed ovaries. +Moquin-Tandon[183] cites from Agardh an instance which seems more +closely to resemble the state of things in the _Baeckea_, and which +occurred in a double hyacinth, wherein both anthers and ovules were +borne on the same placenta. Probably, though the fact is not stated, the +ovary of the hyacinth was open; and we are told that the flower was +double--that it was, in fact, modified and changed in more organs than +one; while in the _Baeckea_ nothing at all unusual was observed till the +ovary was cut open. The style was present even in those flowers where +there was no axile placenta; hence in these cases it could not be, as +Lindley stated it to be in the closely allied _Babingtonia_, a +prolongation of the placenta.[184] + +=Formation of pollen within the ovules.=--This has now been recorded in +two instances by Mr. S. J. A. Salter in _Passiflora caerulea_ and in _P. +palmata_,[185] and by the author in _Rosa arvensis_.[186] + +[Illustration: FIG. 99.--Pollen within the ovule of _Passiflora_ (after +Salter).] + +In the case of the passion-flower there were various malformations in +the ovaries, which were all more or less split open at the distal end, +indicating a tendency towards dialysis. The pollen-bearing ovules were +borne on the edges of these ovaries, and presented various intermediate +conditions between anthers and ovules, commencing at the distal +extremity of the carpel with a bi-lobed anther, and passing in series to +the base of the ovary, an antheroid body of ovule-like form, a modified +ovule containing pollen, an ovule departing from a perfectly natural +condition only in the development of a few grains of pollen in its +nucleus, and, finally, a perfect, normal ovule. + +In the flowers of the Rose the stamens exhibited almost every +conceivable gradation between their ordinary form and that of the +carpels, while some of the ovules contained pollen in greater or less +abundance. Speaking generally, the most common state of things in these +flowers was the occurrence on the throat of the calyx, in the position +ordinarily occupied by the stamens, and sometimes mingled with those +organs, of twisted, ribbon-like filaments, which bore about the centre +one or more pendulous, anatropous ovules on their margins. Immediately +above the latter organs were the anther-lobes, more or less perfectly +developed, and surmounting these a long style, terminating in a fringed, +funnel-shaped stigma. Sometimes the ovules were perfect, at other times +the nucleus protruded through the foramen, while in a third set the +nucleus was included within the tegument, the ovules having in all +respects their natural external conformation, containing, however, not +only pollen-grains, but also a layer of those peculiar spheroidal cells, +including a fibrous deposit, which are among the normal constituents of +the anther. In one case, where the coat of the ovule was imperfect, and +allowed the nucleus to protrude, the pollen was evidently contained +within the central mass of the structure. In this instance the fibrous +cells were not detected, these being only found in cases where the +investment of the ovule was perfect; and hence it seems likely that the +fibrous cells were part of the coat of the ovule, while the pollen was +formed within the nucleus. In no case was any trace of embryo sac to be +seen. + +The main interest, as Mr. Salter remarks, in these cases is +physiological; so far as structure alone is concerned, there does not +appear any reason why pollen-grains should not be developed in any +portion of the plant; the mother cells in which the pollen is formed not +differing, to all outward appearance, from any other cells, unless it be +in size. + +The fundamental unity of construction in all the organs of plants could +hardly be better illustrated than by these cases; while, in spite of +their exceptional nature, they must be of great interest +physiologically, as showing the wide limits of possible variation which +thus may even involve the sex, "for an ovule to develop pollen within +its interior," says Mr. Salter, "is equivalent to an ovum in an animal +being converted into a capsule of spermatozoa. It is a conversion of +germ into sperm, the most complete violation of individuality and unity +of sex. * * * * The occurrence of an antheroid ovule and a normal ovule +on the same carpellary leaf realises the simplest and the most absolute +form of hermaphroditism." + +It must, however, be remarked that the term substitution would be +preferable to conversion. There is, at present, no evidence to show that +the germinal vesicles were present in these cases; on the other hand, it +seems most probable that they were not, so that the presence of the +pollen-cells must be considered as simply adventitious. It can hardly be +that they were, in the first instance, germinal vesicles, which, in +course of time, became so modified as to assume the appearance of +pollen-grains. Between the nucleus of the ovule and the tubercle of +cellular tissue constituting the primordial anther, there is little or +no difference, so that it may be said that, for a time, there is no +distinction of sex in the nascent flower, but as development goes on, +the difference becomes perceptible. It cannot at present be stated what +precise circumstances induce the one mass to form mother-cells and +pollen-grains, and the other to develop an embryo sac and germinal +vesicles. Position and external circumstances may have some indirect +effect, and it may, perhaps, be significant that in all the instances of +polliniferous ovules, the ovular structures have been exposed on an open +carpel or otherwise, in place of being confined within the cavity of a +closed ovary, as under ordinary circumstances. Even among Conifers the +ovuligerous scales are so closely packed that there is little or no +exposure of the ovules. But, apart from all speculative notions as to +the relation between the structure and functions of the anther and of +the ovule respectively, and of the possibility or the reverse of +parthenogenesis, it will clearly be necessary in any future alleged +occurrence of the latter phenomenon to ascertain whether any or all of +the apparent ovules are, or are not, anthers in disguise. + +=Homomorphic flowers of "Compositae."=--In a large section of the +_Compositae_ there is, as is well known, a distinction between the +florets of the "disc" and those of the "ray," the latter being ligulate, +the former tubular. + +In what are erroneously called double flowers in this order, _e.g._ in +the Chrysanthemum, Dahlia, &c. &c., the florets are all ligulate. This +change is sometimes classed with peloria, but there is no abnormal +regularity in these cases. On the other hand, were the ligulate florets +to be all replaced by tubular ones, the term peloria would be more +strictly applicable. It will be remembered that in the sub-order +_Liguliflorae_, the florets are naturally all ligulate, so that the +change above mentioned is not in itself a very grave one. + +=Heterotaxy affecting the inflorescence.=--Under the head of +Prolification, Heterogamy, &c., various deviations from the normal +inflorescence are alluded to. In this place, therefore, it is only +necessary to mention certain rare deviations from the customary +arrangement of the inflorescence, such as the change from a definite +centrifugal form of inflorescence to an indefinite centripetal one. This +occurs occasionally in roses, where the shoot, instead of terminating in +a flower-bud, lengthens and bears the flower-bud on its sides as in a +raceme. + +In the hyacinth, the inflorescence of which is properly indefinite, the +terminal flower may frequently be found to expand first, though in order +of development it may have been the last formed. + +It occasionally happens that certain plants will, contrary to their +usual custom, bloom twice in the same season; this usually arises from +the premature development of buds which, under ordinary circumstances, +would not unfold till the following spring. In these instances of what +the French term "fleuraison anticipee," the position of inflorescence is +not changed, but there are other cases where the position of the +inflorescence is altered, as in the laburnum, where, in some seasons, +racemes may be seen springing from short lateral "spurs" along the sides +of the branches, as well as from the extremities of long shoots. + +Of a similar nature are those cases wherein stems or branches usually +sterile become fertile; this happens in _Equisetaceae_,[187] in +_Restiaceae_, and other orders. In the equisetums, the condition in +question has been specially noticed to occur after prolonged drought. + +_Equisetaceae_ are likewise subject to an anomaly called by Duval Jouve +interruption of the spike, and wherein the scales bearing the spore +cases are separated by whorls of branches instead of forming one compact +unbroken spike as usual. + +This alternation of the organs of vegetation and reproduction may also +be seen occasionally in _Typha_, and other plants. + +Kirschleger describes a case in which the male catkins of _Salix +cinerea_ were placed at the ends of the branches instead of being +lateral productions; moreover the usual articulation was not formed, so +that the catkin was persistent instead of deciduous.[188] + +=Supra-soriferous ferns.=--In the great majority of ferns the sori or +clusters of spore cases are placed on the under surface of the fronds; +nevertheless, a few cases are on record where the fructification is +produced on the upper as well as on the lower surface, and sometimes +abundantly so. This occasionally happens from the elongation of the +normally placed sorus, which thus extends to the margin, and returns on +the upper side, when the sori chance to be placed opposite to the +marginal crenatures. But it is also frequently the case that the sori +are produced on the upper side, distinctly within the margin, and where +there are no corresponding sori beneath. Those varieties which have the +margin crenated or lobed seem most liable to assume this abnormal +supra-soriferous condition. Among the ferns in which this condition has +been observed are the following: _Scolopendrium vulgare_, _Polypodium +anomalum_, Hook., _Asplenium Trichomanes_, _Cionidium Moorei_.[189] + +FOOTNOTES: + +[147] 'Gard. Chron.' 1852, p. 51. + +[148] 'Flora.' 1858, pp. 32-42. + +[149] 'Journ. Linn. Soc.,' vol. vi; "Botany," 1862, p. 24. + +[150] 'Tijdschrift voor Natuur. Geschied,' 1836, vol. iii, tab. vii, p. +171. + +[151] Roeper, 'Enum. Euphorb.,' p. 19. Bernhardi, 'Linnaea,' vii, p. +561, tab. xiv, f. 1. Wydler, "Subcotyled. sprossbildung," 'Flora,' 1850, +p. 337. Hooker, 'Trans. Linn. Soc.,' vol. xxiv, p. 20 (_Welwitschia_). + +[152] 'Misc. Austriac. ad Bot.,' vol. i, p. 133, t. 5. + +[153] See also Carriere, 'Revue Horticole,' 1866, p. 442; and as to +pears, Radlkofer in 'Bericht ueber die Thaetigkert der Baierischen +Gartenbau Gesellschaft,' 1862, p. 74, t. i. + +[154] 'Flora Indica,' p. 23. + +[155] 'Bull. Soc. Bot. Fr.,' 1856, p. 53. + +[156] 'Trans. Linn. Soc.' xxvi, p. 142, tab. iv, B. + +[157] "Si arbusculam, quae in olla antea posita, quotannis floruit et +fructus protulit, deinde deponamus in uberiori terra calidi caldarii, +proferet illa per plures annos multos ac frondosos ramos, sine ullo +fructu. Id quod argumento est, folia inde crescere, unde prius enati +sunt flores; quemadmodum vicissim, quod in folia nunc succrescit, id, +natura ita moderante, in flores mutatur, si eadem arbor iterum in olla +seritur."--Linnaeus, 'Prolepsis,' Sec. iii. + +[158] 'Rev. Hortic.' May, 1868, 'Gardeners' Chronicle,' 1868, pp. 572, +737. + +[159] Cited in 'Annals Nat. Hist.,' 1845, vol. xv, p. 177. + +[160] 'Ann. Scienc. Nat.,' vol. xiv, 1860, p. 13. + +[161] Naudin, 'Ann. Sc. Nat.,' 2nd ser., 1840, vol. xiv, p. 14, fig. 6, +pl. i (_Drosera_). St. Hilaire, 'Comptes Rendus,' ix, p. 437. + +[162] Hance, 'Hook. Journ. Botany,' 1849, vol. i, p. 141, pl. v. + +[163] Booth, 'Gard. Chron.,' Jan. 1st, 1853, p. 4. + +[164] Lindley, 'Theory of Horticulture,' ed. 2, p. 273. + +[165] 'Hook. Journ. of Botany,' 1852, iv, p. 206. See also the curious +_Begonia gemmipara_, 'Hook. fil. Illust. Himal. Plant.,' t. xiv. + +[166] 'Phytanth.,' n. 36, _d._ + +[167] 'Ann. Scienc. Nat.,' 3rd series. 1853. vol. xix, p. 251, tab. 14. + +[168] Carriere, 'Revue Horticole.' 1868, p. 184. + +[169] 'Gard. Chron.,' 1858, p. 556. + +[170] 1863, p. 556, &c. + +[171] 'Ann. Nat. Hist.,' 1845, vol. xvi, p. 355. + +[172] See also Lindley, 'Veg. Kingd.,' p. 109 et 116_a_, where the views +of Raspail, R. Brown, Mohl, Henslow, and others, are discussed. + +[173] It has been observed that if a plant is supplied with copious +nourishment the flowering-period is delayed; but that moderate or even +scanty nourishment accelerates it. Goethe, 'Metam.,' Sec. 30. See also +Wolff, 'Theoria Generationis,' 1759; Linn. 'Prolepsis,' Secs. 3 and 10. + +[174] Moquin-Tandon, p. 384; also Lindl., 'Elements of Botany,' p. 65, +fig. 130; "Theory of Horticulture," p. 86. 'Gard. Chron.,' 1851, p. 723; +Irmish, 'Flora,' 1858, p. 38, &c. + +[175] Caspary, 'Bull. Soc. Bot. Fr.,' vol. vi, 1859, p. 235; also Payer, +ibid., vol. i, 1854. p. 283. + +[176] Trecul, 'Ann. Sc. Nat.,' 2nd ser., vol. xx, p. 339. + +[177] 'Bull. Soc. Bot. Fr.,' vol. vii, 1858, p. 331. + +[178] 'Bull. Soc. Bot. Fr.,' vol. i. p. 306, vol. v, p. 115. 'Illustr. +Hortic.,' xii, 1865, Misc. 79. 'Rev. Horticole,' 1860 p. 204, et 1867 p. +43. + +[179] 'Bull. Soc. Bot. Fr.,' 1858, p. 685. + +[180] The structure of this flower is discussed at some length in a +paper by the author on axillary prolification. 'Trans. Linn. Soc.,' vol. +xxiii, p. 486, t. liv. fig. 3. See also 'Clos. Bull. Soc. Bot. Fr.,' +vol. v, 1855, p. 672. Seringe et Heyland, 'Bull. Bot.,' i, p. 8. 'Pallas +Enum. Plant. Hort. Demidoff,' append, c, ic. + +[181] 'Adansonia,' i, 181. + +[182] 'Adansonia.' vol. iii, p. 351, tab. xii. + +[183] 'Elem. Terat. Veget.,' p, 218. + +[184] Masters, 'Journ. Linn. Soc.,' vol. ix, 1866, p. 334. + +[185] 'Trans. Linn. Soc.,' vol. xxiv, p. 143. tab. xxiv. + +[186] 'Brit. Assoc. Report,' Dundee, 1867; and Seemann's 'Journal of +Botany,' 1867, p. 319, tab. lxxii, figs. B 1-9. + +[187] Duval Jouve, 'Hist. Equiset. France.' 1864, p. 154. + +[188] 'Flora,' t. xxiv, 1841, p. 340. + +[189] Moore, 'Nature-Printed British Ferns,' 8vo edition, vol. ii. p. +135. tab. lxxxv, B, &c. + + + + +CHAPTER IV. + +HETEROGAMY. + + +This term is here intended to apply to all those cases in which the +arrangement of the sexual organs is different from what it is +habitually. It is evident that in many instances there is no +malformation, no monstrosity, but rather a restoration of organs +habitually suppressed, a tendency towards structural completeness +rather than the reverse. It must be also understood that the following +remarks apply to structural points only, and are not intended to include +the question of function. The occurrence of heteromorphic unions renders +it necessary to keep in mind that plants hermaphrodite as to structure +are by no means necessarily so as to function. + +The simplest case of this alteration in the relative position of the +sexes is that which occurs in monoecious plants, where the male and +female flowers have a definite position, but which in exceptional +instances is altered. + +=Change in the relative position of male and female flowers= may thus +occur in any monoecious plant. Cultivated maize, _Zea Mays_, +frequently exhibits alterations of this kind; under ordinary +circumstances, the male inflorescence is a compound spike, occupying the +extremity of the stem, while the female flowers are borne in simple +spikes at a lower level, but specimens may now and then be found where +the sexes are mixed in the same inflorescence; the upper branching +panicle usually containing male flowers only, under these circumstances, +bears female flowers also.[190] In like manner, but less frequently, the +female inflorescence occasionally produces male flowers as well. + +Among the species of _Carex_ it is a common thing for the terminal spike +to consist of male flowers at the top, and female flowers at the base; +the converse of this, where the female flowers are at the summit of the +spike, is much more uncommon. An illustration of this occurrence is +given in the figure (fig. 100). Among the _Coniferae_ numerous instances +have been recorded of the presence of male and female flowers on the +same spike, thus Mr. now Professor Alexander Dickson exhibited at the +Botanical Society of Edinburgh in July, 1860, some malformed cones of +_Abies excelsa_, in which the inferior part of the axis was covered with +stamens, whilst the terminal portion produced bracts and scales like an +ordinary female cone. The stamens of the lower division were serially +continuous with the bracts above. Some of the lower scales of the female +portion were in the axils of the uppermost stamens, which last were +somewhat modified, the anther cells being diminished, whilst the +scale-like crest had become more elongated and pointed, in fact, more or +less resembling the ordinary bracts.[191] Mohl, Schleiden, and A. Braun +have observed similar cones in _Pinus alba_, and Cramer figures and +describes androgynous cones in _Larix microcarpa_. C. A. Meyer ('Bull. +Phys. Math.,' t. x, 1850) also describes some catkins of _Alnus +fruticosa_ which bore male flowers at the top, and female flowers at the +base. + +[Illustration: FIG. 100.--Spike of _Carex acuta_, with female flowers at +the summit.] + + On the subject of this section the reader may consult A. + Braun,. 'Das Individ.,' 1853, p. 65. Caspary, 'De Abietin. + flor. fem. struct. morphol.' Schleiden. 'Principles,' English + edition, p. 299. Mohl, 'Verm. Schrift.,' p. 45. Meyen in + 'Wiegm. Archiv.,' 1838, p. 155. Cramer, 'Bildungsabweich,' p. + 4, tab. v, figs. 13-17. Parlatore, 'Ann. Sc. Nat.,' ser. iv, + vol. xvi, p. 215, tab. 13A. See also under the head of + Prolification, Substitutions, &c. + +=Change from the monoecious to the dioecious condition.=--This is of +less frequent occurrence than might have been anticipated. In the +'Gardeners' Chronicle,' 1847, pp. 541 and 558, several instances are +noted of walnut trees bearing female flowers to the exclusion of males. +The mulberry tree has also been noticed to produce female blossoms only, +while in other plants male flowers only are developed. + +It seems probable that the age of the plant may have something to do +with this production of flowers of one sex to the exclusion of the +other. + +=Change from the dioecious to the monoecious condition.--Androgynism.=-- +This is of far more common occurrence than the preceding. + +[Illustration: FIG. 101.--Monoecious inflorescence of Hop.] + +In the hop (_Humulus Lupulus_), when monoecious, the female catkins +are usually borne on the ends of the branches as shown in the cut (fig. +101), and a similar thing has been noticed in _Urtica dioica_ by Clos, +'Bull. Soc. Bot. France,' vol. 9, p. 7. + +Baillon ('Etudes du groupe des Euphorbiacees,' p. 205) mentions the +following species of that order as having been seen by him with +monoecious inflorescence: _Schismatopera distichophylla_, _Mozinna +peltata_, _Hermesia castaneifolia_. Oliver mentions ('Hook. Icon. +Plant.,' t. 1044) that in _Leitneria floridana_ the upper scales of the +male catkin occasionally subtend an ovary. + +It would seem that external conditions have some effect in determining +the formation of one sex, as in some species of _Carex_, while in the +case of _Salix repens_, Hampe[192] says that when grown partially or for +a time under water, those twigs which are thrust up above the surface +bear female flowers, while those twigs that blossom after the water is +dried up, produce male flowers only. + +Carriere[193] says that a plant of _Stauntonia latifolia_ which for some +years produced stamens only, now produces flowers of both sexes; it was +dioecious, but is now monoecious. The same author alludes to a +similar occurrence in _Juniperus Virginiana_. The hops is also said to +vary in sexual characteristics from time to time.[194] In addition to +the genera, already named, in which this production of flowers of both +sexes has been observed may be mentioned _Taxus! Gunnera! Urtica! +Mercurialis! Restio! Cannabis! Salix! Humulus!_ as well as others in +which the change is less frequent. + +Among cryptogams a similar change occurs. As an illustration may be +cited _Leucobryum giganteum_, as quoted from Mueller in Henfrey's +'Botanical Gazette,' i, p. 100. + + As to androgynous willows, in addition to the references given + under the head of Substitution of stamens for pistils, see + Schlechtendal, 'Flora Berol.,' ii. p. 259. Tausch, 'Bot. + Zeit.,' 1833, i. p. 229. Koch, 'Synops. Flor. Germ.,' 740. + Host, 'Flor. Aust,.' ii, p. 641 (_S. mirabilis_). See also + Hegelmaier, 'Wuerttemberg Naturwissenshaft Jahreshefte,' 1866, + p. 30. Other references to less accessible works are given in + 'Linnaea,' xiv, p. 372. + +=Change from hermaphroditism to unisexuality.=--Many flowers ordinarily +hermaphrodite as to structure, become unisexual by the abortion or +suppression of their stamens, or of their carpels, as the case may be. +This phenomenon is lessened in interest since the demonstration of the +fact by Darwin and others, that many plants, structurally hermaphrodite, +require for the full and perfect performance of their functions the +cooperation of the stamens and pistils, belonging to different +individuals of the same species. + +Some of the _Ranunculaceae_ constantly exhibit a tendency towards the +dioecious condition, and the rarity with which perfect seeds of +_Ranunculus Ficaria_ are formed is to be attributed, in great measure, +to the deficiency of pollen in the anthers of these flowers. _Ranunculus +auricomus_ also is frequently sterile. Specimens of _Ranunculus +bulbosus_ may be met with in which every flower is furnished with +carpels, most of which have evidently been fertilised, although there +are no perfect stamens in the flowers. + +Knight and other vegetable physiologists have been of opinion that a +high temperature favours the production of stamens, while a lower degree +of heat is considered more favorable to the production of pistils, and +in this way the occurrence of "blind" strawberries has been accounted +for. Mr. R. Thompson, writing on this subject, speaks of a plantation of +Hautbois strawberries which in one season were wholly sterile, and +accounts for the circumstance as follows: the plants were taken from the +bearing beds the year previous, and were planted in a rich well-manured +border, in which they started rapidly into too great luxuriance, the +growth being to leaves rather than to fruit. The following season these +same plants bore a most abundant crop, hence these plants were +accidentally prevented from perfecting their female organs.[195] + +Mr. Darwin[196] cites from various sources the following details +relating to strawberries which it may be useful to insert in this place, +as throwing some light upon the production of unisexual flowers. +"Several English varieties, which in this country are free from any such +tendency, when cultivated in rich soils under the climate of North +America commonly produce plants with separate sexes. Thus, a whole acre +of Keen's seedlings in the United States has been observed to be almost +sterile in the absence of male flowers; but the more general rule is, +that the male plants over-run the females.... The most successful +cultivators in Ohio plant, for every seven rows of pistillate flowers, +one row of hermaphrodites, which afford pollen for both kinds; but the +hermaphrodites, owing to their expenditure in the production of pollen, +bear less fruit than the female plants." + +_Stratiotes aloides_ has been said to produce its carpels with greater +abundance towards the northern limits of its geographical distribution, +and its stamens, on the other hand, are stated to be more frequently +developed in more southern districts. + +_Honckenya peploides_ affords another illustration of the sexual +arrangements in the flower being altered as it would seem by climatal +conditions. Thus, in the United States, according to Professor Asa Gray, +the flowers are frequently hermaphrodite, while in this country they are +usually sub-dioecious.[197] + +Treviranus[198] says that the flowers of _Hippuris_ and _Callitriche_ +are apt to be hermaphrodite in summer, but female only at a later +period. + +For further remarks on this subject, see sections relating to +suppression of stamens and pistils. + +=Change from unisexuality to hermaphroditism.=--This occurrence depends +on one of two causes, either organs are developed (stamens or pistils as +the case may be), which are habitually absent in the particular flower; +or some of the stamens may be more or less completely converted into or +replaced by pistils, or _vice versa_. + +The first condition is the opposite of suppression; it is, as it were, a +restoration of symmetry, and might be included under the head of regular +peloria, inasmuch as certain organs which habitually undergo suppression +at a certain stage in their development, by exception, go on growing, +and produce a perfect, instead of an imperfect flower. In teratological +records it is not always stated clearly to which of the two above-named +causes the unusual hermaphroditism belongs, though it is generally easy +to ascertain this point. Very many, perhaps all, diclinous flowers may, +under certain conditions, become perfect, at least structurally. I have +myself seen hermaphrodite flowers in _Cucurbita_,[199] _Mercurialis_, +_Cannabis_, _Zea Mays_, and _Aucuba japonica_, as well as in many +_Restiaceae_, notably _Cannamois virgata_ and _Lepyrodia hermaphrodita_. +_Spinacia oleracea_, _Rhodiola rosea_, _Cachrys taurica_, and _Empetrum +nigrum_ are also occasionally hermaphrodite. + +Gubler[200] alludes to a similar occurrence in _Pistacia Lentiscus_, +wherein, however, he adds that there was a deficiency of pollen in the +flowers. + +Schnizlein[201] observed hermaphrodite flowers in the beech, _Fagus +sylvatica_, the ovaries being smaller than usual, and the stamens +epigynous. + +Baillon[202] enumerates the following _Euphorbiaceae_ as having +exceptionally produced hermaphrodite flowers, _Crozophora tinctoria_, +_Suregada_ sp., _Phyllanthus longifolius_, _Breynia_ sp., _Philyra +brasiliensis_, _Ricinus communis_, _Conceveiba macrophylla_, _Cluytia +semperflorens_, _Wall_, non _Roxb_. _Mercurialis annua_ and +_Cleistanthus polystachyus_. + +In some of these cases the hermaphroditism is due to the development of +anthers on the usually barren staminodes, though, in other cases, the +stamens would seem to be separate, independent formations, as they do +not occupy the same relative position that the ordinary stamens would do +if developed.[203] + +[Illustration: FIG. 102.--Flower of _Fuchsia_ in which the calyx was +leafy, the petals normal (reflexed in the figure), the stamens partially +converted into ovaries, the ordinary inferior ovary being absent. See +Substitution.] + +Robert Brown[204] observed stamens within the utricle of _Carex acuta_, +and Gay is stated by Moquin ('El. Ter. Veg.,' p. 343) to have observed +a similar occurrence in _Carex glauca_. + +Paasch[205] observed a similar occurrence in _C. caespitosa_, and +Schauer, in _C. paludosa_,[206] though in the latter instance the case +seems to have been one of transformation or substitution rather than one +of hermaphroditism. + +The second cause of this pseudo-hermaphroditism is due either to the +more or less perfect mutation of male and female organs, or it may be to +the complete absence of one and its replacement by another, as when out +of many stamens, one or more are deficient, and their places occupied by +carpels. This happens very frequently in willows and poplars, and has +been seen in the beech.[207] + +[Illustration: FIG. 103.--Hermaphrodite flower of _Carica Papaya_.] + +In _Begonia frigida_[208] the anomaly is increased by the position of +the ovaries above, the perianth, a position due, not to any solution or +detachment of the latter from the former, but simply to the presence of +ovaries where, under ordinary circumstances, stamens only are formed, +as happened also in a garden variety of a _Fuchsia_, wherein, however, +the change was less perfect than in the _Begonia_, and in which, as the +flower is naturally hermaphrodite, the alteration is of the less +importance. + +[Illustration FIG. 104.--Ovuliferous anthers--_Cucurbita_.] + +In hermaphrodite flowers of _Carica Papaya_ (fig. 103) there is a single +row of five stamens instead of two rows of five each as in the normal +male flowers, the position of the second or inner row of stamens being +occupied by five carpels, which, however, are not adherent to the +corolla as the stamens are, thus, supposing the arrangement of parts in +the normal male flowers to be as follows: + + --------------------------- + s s s s s + --------------------------- + | p p p p p + | st st st st st + | st st st st st + | + +That of the hermaphrodite blossoms would be, in brief, as follows: + + | 5 s + |------------ + | 5 p + | 5 st + | 5 c + | + +One of the most curious cases of this kind recorded is one mentioned by +Mr. Berkeley,[209] wherein a large white-seeded gourd presented a +majority of flowers in which the pollen was replaced by ovules. It would +seem probable from the appearances presented by the figure that these +ovules were, some of them, polliniferous, like those of the +_Passiflora_, &c., described at p. 185, but nothing is stated on the +subject. + +See also section on Regular Peloria, Substitution, Pistillody of the +stamens, &c. + +FOOTNOTES: + +[190] See also Clos., 'Mem. Acad. Toulouse,' sixth ser., t. iii, pp. +294-305. Scott, 'Trans. Bot. Soc. Edinburgh,' t. viii, p. 60. Wigand, +'Flora,' 1856, p. 707. + +[191] Professor Dickson concludes from the examination of these +structures that the male cone, consisting of simple stamens developed on +one common axis, must be regarded as a simple male flower, while the +axillary scales of the female cone are by him compared with the +flattened shoots of _Ruscus_. + +[192] 'Linnaea,' xiv, 367. + +[193] Rev. Hortic.,' January, 1867. + +[194] See Royle, 'Man. Materia Medica,' ed. 1, p. 567. + +[195] Thomson, 'Gardener's Assistant,' p. 577. + +[196] 'Variation of Animals and Plants,' i, 353. + +[197] Babington, 'Ann. Nat. Hist.,' vol. ix, 1852, p. 156. + +[198] 'Phys. der Gewaechse,' ii, p. 323. + +[199] See also Schlechtendal, 'Linnaea,' viii, p. 623, and Lindley, 'Veg. +Kingd.,' p. 315. + +[200] 'Bull. Soc. Bot. France,' vol. ix, p. 81. + +[201] Cited in Henfrey, 'Bot. Gazette.' 3, p. 11. + +[202] Baillon. 'Etudes du Groupe des Euphorbiacees,' p. 205, tab. xv, +fig. 19, tab. xix, fig. 31. + +[203] See also Guillemin, 'Mem. Soc. Nat. Hist. Paris,' I, p. 16; +hermaphrodite flowers in _Euphorbia esula_. + +[204] 'Prod. Flor. N. Holl.,' p. 242. + +[205] 'Bot. Zeit.,' 1837, p. 335. + +[206] 'Pflanz, Terat.,' von Moquin-Tandon, p. 208. + +[207] Schnizlein, loc. cit. + +[208] 'Bot. Mag.,' tab. 5160, fig. 4. See also 'Gard. Chron.,' 1860, pp. +146, 170; 1861, p. 1092. + +[209] 'Gard. Chron.,' 1851, p. 499. + + + + +CHAPTER V. + +ALTERATIONS IN THE DIRECTION OF ORGANS. + + +The deviations from the ordinary direction of organs partake for the +most part more of the nature of variations than of absolute malposition +or displacement. It must also be borne in mind how frequently the +direction of the leaves, or of the flower, varies according to the +stage of development which it has arrived at, to unequal or +disproportionate growth of some parts, or to the presence of some +impediment either accidental or resulting from the natural growth of the +plant. These and other causes tend to alter the direction of parts very +materially. + +=Change in the direction of axile organs, roots, stems, &c.=--The roots +frequently exhibit good illustrations of the effect of the causes above +mentioned in altering the natural direction. The roots are put out of +their course by meeting with any obstacle in their way. Almost the only +exception to the rule in accordance with which roots descend under +natural circumstances, is that furnished by _Trapa natans_, the roots of +which in germination are directed upwards towards the surface of the +water. So in _Sechium edule_, the seed of which germinates while still +in the fruit, the roots are necessarily, owing to the inverted position +of the embryo, directed upwards in the first instance. + +A downward direction of the stem or branches occurs in many weak-stemmed +plants growing upon rocks or walls, or in trees with very long slender +branches as in _Salix Babylonica_, and the condition may often be +produced artificially as in the weeping ash. + +The opposite change occurs in what are termed fastigiate varieties, +where the branches, in place of assuming more or less of a horizontal +direction, become erect and nearly parallel with the main stem as in the +Lombardy poplar, which is supposed to be merely a form of the black +Italian poplar. + +M. de Selys-Longchamps has described a similar occurrence in another +species of Poplar (_P. virginiana_ Desf.), and amongst a number of +seedling plants fastigiate varieties may frequently be found, which may +be perpetuated by cuttings or grafts, or sometimes even by seed; hence +the origin of fastigiate varieties of elms, oaks, thorns, chestnuts, and +other plants which may be met with in the nurseries. + +Sometimes when the top of the main stem is destroyed by disease or +accident, one of the heretofore lateral shoots takes its place, and +continues the development of the tree in the original direction. It is +often an object with the gardener to restore the symmetry of an injured +tree so that its beauty may ultimately not be impaired.[210] + +Climate appears sometimes to have some influence on the direction of +branches, thus Dr. Falconer, as quoted by Darwin,[211] relates that in +the hotter parts of India "the English Ribston-pippin apple, a Himalayan +oak, a Prunus and a Pyrus all assume a fastigiate or pyramidal habit, +and this fact is the more interesting as a Chinese tropical species of +_Pyrus_ naturally has this habit of growth. Nevertheless many of the +fastigiate varieties seen in gardens have originated in this country by +variation of seeds or buds." + +M. Carriere has also recorded a curious circumstance with reference to +the fastigiate variety of the false acacia _Robinia pseudacacia_; he +states that if a cutting or a graft be taken from the upper portion of +the tree, the fastigiate habit will be reproduced, and the branches will +be furrowed and covered with short prickles; but if the plant be +multiplied by detaching portions of the root-stock, then instead of +getting a pyramidal tree with erect branches, a spreading bushy shrub is +produced, with more or less horizontal, cylindrical branches, destitute +of prickles.[212] + +=Eversion of the axis.=--In the case of the fig, the peculiar +inflorescence is usually explained on the supposition that the +termination of the axis becomes concave, during growth, bearing the true +flowers in the hollow thus formed. The cavity in this case would +probably be due not to any real process of excavation, but to a +disproportionate growth of the outer as contrasted with the central +parts of the fig. Some species of _Sempervivum_ have a similar mode of +growth, so that ultimately a kind of tube is formed, lined by the +leaves, the central and innermost being the youngest. The hip of the +Rose may be explained in a similar manner by the greater proportionate +growth of the outer as contrasted with the central portions of the apex +of the flower-stalk. In cases of median prolification, already referred +to, the process is reversed, the central portions then elongate into a +shoot and no cavity is formed. A fig observed by Zuccarini (figs. 105, +106) appears to have been formed in a similar manner, the flower-bearing +summit of the stalk not being contracted as usual, the flowers projected +beyond the orifice of the fig. If this view be correct the case would be +one rather of lengthening of the axis than of absolute eversion since it +was never inverted. + +[Illustration: FIG. 105.--Fig showing prolonged inflorescence and +projecting flowers.] + +[Illustration: FIG. 106.--Section of the same.] + +=Altered direction of leaves.=--The leaves partake more or less of the +altered direction of the axis, as in fastigiate elms, but this is not +universally the case, for though the stem is bent downwards the leaves +may be placed in the opposite direction; thus in some specimens of +_Galium Aparine_ growing on the side of a cliff from which there had +been a fall of chalk, the stems, owing apparently to the landslip, were +pendent, but the leaves were abruptly bent upwards. + +One of the most singular instances of an inverted direction of the +leaves is that presented by a turnip (fig. 107) presented to the Museum +of King's College, London, by the late Professor Edward Forbes. The +turnip is hollow in the interior and the majority of the leaves +springing from its apex instead of ascending into the light and air +become bent downwards so as to occupy the cavity, and in such a manner +as to bring to mind the position of an inverted embryo in a seed. + +[Illustration: FIG. 107.--Hollow turnip, showing some of the leaves +inverted and occupying the cavity.] + +=Altered direction of the flower and its parts.=--The changes which take +place in the relative position either of the flower as a whole or of its +several parts during growth are well known, as also are the relations +which some of these movements bear to the process of fertilisation, so +that but little space need here be given to the subject beyond what is +necessary to point out the frequent changes of direction which +necessarily accompany various deviations from the ordinary form and +arrangement of parts. + +In cases where an habitually irregular flower becomes regular, the +change in form is frequently associated with an alteration in direction +both of the flower as a whole and, to a greater or less extent, of its +individual members, for instance of _Gloxinia_, the normal flowers of +which are irregular and pendent, there is now in common cultivation a +peloriate race in which the flowers are regular in form and erect in +position. + +[Illustration: FIG. 108.--Flower of normal _Gloxinia_.] + +[Illustration: FIG. 109.--Flower of _Gloxinia_, erect and regular +(regular _Peloria_).] + +Fig. 108 shows the usual irregular form of _Gloxinia_, with which may be +contrasted figs. 109, 110 and 111. + +Fig. 109 shows the regular erect form; fig. 110 the calyx of the same +flower; while in fig. 111 are shown the stamens and style of the two +plants respectively. In the upper figure the style of the peloriate +variety is shown as nearly straight, and the stamens undergo a +corresponding change. No doubt the relative fertility and capacity for +impregnation of the two varieties is affected in proportion to the +change of form. The Gloxinia affords an instance of regular congenital +peloria in which the regularity of form and the erect direction are due +to an arrest, not of growth, but of development, in consequence of which +the changes that ordinarily ensue during the progress of the flower from +its juvenile to its fully formed condition do not take place. + +[Illustration: FIG. 110.--Calyx of erect _Gloxinia_.] + +[Illustration: FIG. 111.--Stamens of erect regular, and of pendent +irregular-flowered _Gloxinia_.] + +A similar alteration accompanies this form of peloria in other flowers +(see Peloria). A change in direction may result also from other +circumstances than those just alluded to. Abortion or suppression of +organs will induce such an alteration; thus in a flower of _Pelargonium_ +now before me three of the five carpels, from some cause or other, are +abortive and much smaller than usual, and the style and the beak-like +torus are bent downwards towards the stunted carpels instead of being, +as they usually are, straight. + +Amongst orchids, where the pedicel of the flower or the ovary is +normally twisted, so that the labellum occupies the anterior or inferior +part of the flower, it frequently happens, in cases of peloria and other +changes, that the primitive position is retained, the twist does not +take place, and so with other resupinate flowers. In Azaleas a curious +deflexion of the parts of the flower may occasionally be met with. Fig. +112 shows an instance of this in which the corolla, the stamens and the +style were abruptly bent downwards: as young flowers of this singular +variety have not been examined it is difficult to form an opinion as to +the cause of this variation. In one plant the change occurred in +connection with the suppression of all the flowers but one in the +cluster, or rather the place of the flowers was occupied by an equal +number of leafy shoots. + +[Illustration: FIG. 112.--Flower of _Azalea_, showing the corolla +reflected.] + +Moquin[213] mentions a flower of _Rosa alpina_ in which two of the +petals were erect, while the remaining ones were much larger and +expanded horizontally. The same author quotes from M. Desmoulins the +case of a species of _Orobanche_, in which a disjunction of the petals +constituting the upper lip took place, thus liberating the style and +allowing it to assume a vertical direction. + +[Illustration: FIG. 113.--Flower of _Cuphea miniata_ enlarged, showing +protrusion and hypertrophy of an erect placenta, after Morren.] + +[Illustration: FIG. 114.--Placenta from the flower shown at fig. 113; +the ovary is membranous and torn, the placenta, erect and ovuliferous, +after Morren.] + +M. Carriere[214] has described an instance wherein two apples were +joined together, a larger and a smaller one; the former was directed +away from the centre of the tree as usual, while the smaller one was +pointed in exactly the opposite direction. The larger fruit had the +customary parchment-like carpels, the smaller was destitute of them. + +Sometimes the direction assumed by one flower as an abnormal occurrence +is the same as that which is proper to an allied species or genus under +natural circumstances; thus flowers of the vine (_Vitis_) have been met +with in which the petals were spreading like a star (_fleurs +avalidouires_), as in the genus _Cissus_.[215] + +Morren describes a curious condition in some flowers of _Cuphea +miniata_, in which the placenta protruded through an orifice in the +ovary, and losing the horizontal direction became erect (figs. 113, +114). A similar occurrence happened in _Lobelia erinus_. To this +condition the Belgian savant gave the name of gymnaxony.[216] + +FOOTNOTES: + +[210] The following details as to the method pursued by Mr. McNab, of +the Edinburgh Botanic Garden, may not be uninteresting in this place. +They are from the pen of Mr. Anderson, and originally appeared in the +'Gardeners' Chronicle.' + +"The mode of inducing leaders to proceed from laterals is a matter of +comparatively little concern among the generality of deciduous trees, +for they are often provided with subsidiary branches around the leader, +at an angle of elevation scarcely less perpendicular, but the laterals +of all Conifers stand, as nearly as possible, at right angles. Imagine +the consternation of most people when the leader of, say, _Picea +nobilis_, _P. Nordmanniana_, or _P. Lowii_ is destroyed." + +In a specimen of the latter plant the leader had been mischievously +destroyed, to remedy which Mr. McNab adopted means which Mr. Anderson +goes on to describe. "Looking from the leader downward to the first tier +of laterals, there appeared to have been a number of adventitious +leaf-buds created, owing to the coronal bud being destroyed. These were +allowed to plump up unmolested until the return of spring, when every +one was scarified or rubbed off but the one nearest the extremity. To +assist its development and restrain the action of the numerous laterals, +every one was cut back in autumn, and this restraint upon the sap acted +so favorably upon the incipient leader as to give it the strength and +stamina of the original leader, so that nothing detrimental was evident +twelve months after the accident had happened, and only a practical eye +could detect that there had been any mishap at all. This beautifully +simple process saved the baby tree. + +"Another example of retrieving lost leaders may be quoted as +illustrative of many in similar circumstances. _Picea Webbiana_ had its +leader completely destroyed down to the first tier of laterals. There +was no such provision left for inducing leaf-buds as was the case with +_P. Lowii_ above referred to. Resort must, therefore, be had to one of +the best favoured laterals, but how is it to be coaxed from the +horizontal position of a lateral to the perpendicular position of a +leader? The uninitiated in these matters, and, in fact, practical +gardeners generally, would at once reply, by supporting to a stake with +the all-powerful Cuba or bast-matting. But no. A far simpler method than +that, namely, by fore-shortening all the laterals of the upper tier but +the one selected for a leader. Nature becomes the handmaid of art here; +for without the slightest prop the lateral gradually raises itself +erect, and takes the place of the lost leader. All that the operator +requires to attend to is the amputation of the laterals until this +adventitious fellow has gained a supremacy. Singular provision in nature +this, which, thanks to the undivided attention of a careful observer, +has been fully appreciated and utilized." + +[211] 'Variation of Animals and Plants,' ii, p. 277. + +[212] Quoted in 'Gard. Chron.,' 1867, p. 654. + +[213] Loc. cit., p. 315. + +[214] 'Rev. Hortic.,' 1868, p. 110. + +[215] Planchon and Mares, 'Ann. Sc. Nat.,' 5 ser., tom. vi, 1866, p. +228, tab. xii. + +[216] 'Bull. Acad. Belg.,' xviii, part ii, p. 293. + + + + +BOOK II. + +DEVIATIONS FROM THE ORDINARY FORM OF ORGANS. + + +In a morphological point of view the form of the various parts or organs +of plants and the changes to which they are subjected during their +development are only second in importance to the diversities of +arrangement and, indeed, in some cases, do not in any degree hold a +second place. + +Taken together, the arrangement, form, and number of the several parts +of the flower, make up what has been termed the symmetry of the +flower.[217] Referring to the assumed standard of comparison, see p. 4, +it will be seen that in the typically regular flower all the various +organs are supposed to be regular in their dimensions and form. At one +time it was even supposed that all flowers, no matter how irregular +they subsequently became, began by being strictly symmetrical or +regular, and that subsequent alterations were produced by inequality of +growth or development. The researches of organogenists have, however, +dispelled this idea of unvarying primordial regularity, by showing that +in many cases flowers are irregular from the very first, that some begin +by being irregular, and subsequently become regular, and even in some +cases resume their original condition during the course of their +development.[218] Under these circumstances an artificial standard of +comparison becomes almost an absolute necessity for the time being. + +Changes of form very generally, but not always, are accompanied with a +change in regularity: thus a flower habitually bi-lateral may assume the +characters of radiating symmetry and _vice versa_. Increase or decrease +of size very frequently also are co-existent with an alteration in the +usual form. + +In the case of the arrangement of organs it is often difficult or +impossible, in the present state of our knowledge, to determine whether +a given arrangement is congenital or acquired subsequently to the first +development, whether for instance an isolation of parts be due to +primordial separation or to a subsequent disunion of originally combined +organs, see p. 58. With reference to the changes in the form of organs, +however, it is in general more easy to ascertain the proximate cause of +the appearance, and thus teratological changes of form may be grouped +according as they are due to, 1, arrest of development; 2, undue or +excessive development; 3, perverted development; and 4, irregular +development; hence the use of the following terms--Stasimorphy, +Pleiomorphy, Metamorphy, and Heteromorphy--to include teratological +changes really or apparently due to one or other of the causes above +mentioned. The classification here adopted is of course to a +considerable extent an arbitrary one and subject to correction or +modification, as the knowledge of the development of the flowers in the +various genera of plants advances. + +FOOTNOTES: + +[217] The word symmetry has been used in very different senses by +different botanists, sometimes as synonymous with "regularity," at other +times to express the assumed typical form of a flower. Payer understands +it to be that arrangement of parts which permits of the whole flower +being divided vertically into two symmetrical halves (bi-lateral +symmetry). Others, again, have applied the term symmetry to the number +of the parts of the flower, reserving the terms "regularity" or +"irregularity" for the form. It is here used in a general sense to +express the plan of the flower, and thus includes the arrangement, form, +and number of its component elements. + +[218] See Baillon, 'Adansonia,' v, 176. + + + + +PART I. + +STASIMORPHY.[219] + + +Deviations from the ordinary form of organs arising from stasis or +arrest of development are included under this heading. + +There are many cases in which the forms proper to a juvenile condition +of the plant are retained for a much longer period than ordinary, or +even throughout the life of the individual growth goes on, but +"development" is checked. Such conditions may even be propagated by seed +or bud. It is a very general thing for botanists to consider these cases +as reversions to a simpler, primitive type, and this may be so; but on +the other hand, they may be degenerations from a complex type, or they +may have no direct relation to any antecedent condition. Stasimorphic +changes affecting principally the relative size of organs--such, for +instance, as the non-development of internodes, or the atrophy or +suppression of parts will be found mentioned in the sections relating to +those subjects. In the present part those alterations which affect the +form of organs principally are treated of. + +FOOTNOTES: + +[219] [Greek: Stasis-morphosis]. + + + + +CHAPTER I. + +PERSISTENCE OF JUVENILE FORMS. + + +The retention in adult life of a form characteristic of an early stage +of development, and therefore usually transient, may be manifested in +any of the organs of the plant. As these cases are for the most part +treated under separate headings, it is here only necessary to allude to +a few, which it is difficult to allocate satisfactorily, while the +reader may be referred for other instances of like nature to the +sections on Peloria, Atrophy, Suppression, Dimorphy, Substitutions, &c. + +[Illustration: FIG. 115.--_Juniperus sinensis_. Two forms of leaves on +branches of the same shrub.] + +=Stasimorphy in the leaves of conifers.=--In many conifers the leaves +produced in the young state of the plant are different, both in +arrangement and form, from those subsequently developed (see pp. 89, +90). But it occasionally happens that the plant continues to form +throughout its existence leaves such as are usually produced only in a +young state; thus M. Gubler ('Bull. Soc. Bot., Fr.,' vol. viii, 1861, p. +527) describes a plant of _Pinus pinea_ in which the primordial, usually +transitory, foliage was permanent, leaves of the ordinary shape not +being developed at all. It more often happens that some only of the +leaves retain their young form while others assume other shapes, see +fig. 115. This happens frequently in the larch and constantly in the +Chinese juniper when it has arrived at a considerable age. In _Cupressus +funebris_ two forms of leaves may often be found on the same plant, the +one representing the juvenile state, the other the more developed +condition. What is very singular, is that a cutting taken from the +branch with leaves of the young form grows up into a shrub bearing +leaves of no other shape, so that an ordinary observer unacquainted with +the history of the plant would imagine that he had to deal with two +distinct species. This fact is the more interesting when compared with +the alternation of generations which takes place among the lower +animals. + +The regular development of all the parts of the flower in a plant +habitually producing irregular flowers is referred to under the head of +Peloria, but it still remains to consider those examples in which some +only of the parts of the flower are affected in this manner.[220] Most +of these cases are elsewhere referred to in this volume under the +particular form of malformation assumed; but the following case may here +be noticed as not coming under any of the previous heads. It is an +instance recorded by Professor Babington ('Phytologist,' August, 1853), +and in which the pod of _Medicago maculata_, which is usually rolled up +like a snail shell and provided with spines, was sickle-shaped and +unarmed. + +FOOTNOTES: + +[220] See a paper of Professor C. Morren's on "Floral Stesomy" in 'Bull. +Acad. Belg.,' t. xix, part ii, p. 519. + + + + +CHAPTER II. + +REGULAR PELORIA. + + +[Illustration: FIG. 116.--Regular Peloria, _Delphinium_.] + +[Illustration: FIG. 117.--Sepal, petal, &c., of regular-flowered +_Delphinium_.] + +[Illustration: FIG. 118--Regular peloria, _Viola_.] + +[Illustration: FIG. 119--Double Violet, flower regular, petals +multiplied, stamens and pistils petaloid.] + +When an habitually irregular flower becomes regular, it does so in one +of two ways; either by the non-development of the irregular portions, or +by the formation of irregular parts in increased number, so that the +symmetry of the flower is rendered perfect, as in the original peloria +of Linnaeus, and which may be called irregular peloria, while the former +case may be called regular peloria. This latter appearance is therefore +congenital, and due to an arrest of development.[221] As the true nature +of these cases has not been in all cases recognised (even Moquin places +them under the head of deformities--they being less entitled to rank in +that class than are the usual flowers), it may be well to cite a few +instances taken from various families. In _Delphinium peregrinum_ I have +met with perfectly regular flowers having five sepals and five oblong +stalked petals, and a similar occurrence has been noted in other +species of this genus. Baillon,[222] in referring to these flowers, +points out the resemblance that they bear to the double varieties of +_Nigella_. In the stellate columbines (_Aquilegia_) of gardens the +tubular petals are replaced by flat ones often in increased numbers. In +violets both forms of peloria occur, that in which there is an unusual +number of spurs, and that in which there are no spurs (var. anectaria). +In the more perfect forms of regular peloria occurring in the last-named +genus the following changes may be noticed: 1, an alteration in the +direction of the flower so that it remains in an erect position, and is +not bent downwards as usual; 2, equality of proportion in the sepals and +petals; 3, absence of spurs, as also of hairs on the lateral petals; 4, +equal stamens whose anthers are sometimes entirely destitute of the +prolonged crest which forms so prominent a feature under ordinary +circumstances; 5, erect, not curved styles, and the stigmas not +prolonged into a beak, but having a more or less capitate form; ovary +with three or five cells, ovules normal. + +These are cases where the change in question is most strongly marked, +the bi-lateral is completely replaced by the radiating symmetry. The +absence of the usual nectary, and of hairs on the side petals, the +alterations in the form of the style, etc., all show how much the +process of fertilisation must be altered from that which occurs under +ordinary circumstances. In some of the double violets now cultivated in +gardens, a similar regularity of proportion in the parts of the flower +may be seen combined with the substitution of petals for stamens and +pistils, and with the development of an increased number of petal-like +organs.[223] Between these cases and the ordinary spurred forms as well +as those with an increased number of spurs, many intermediate forms may +be met with. That such regularity should occur in this family is not to +be wondered at seeing that there is a whole subdivision of the order +(_Alsodeiae_) in which regular flowers are the rule. + +In cultivated Pelargoniums the central flower of the umbel or "truss" +frequently retains its regularity of proportion, so as closely to +approximate to the normal condition in the allied genus _Geranium_; this +resemblance is rendered greater by the fact that, under such +circumstances, the patches of darker colour characteristic of the +ordinary flower are completely wanting; the flower is as uniform in +colour as in shape. Even the nectary which is adherent to the upper +surface of the pedicel in the normal flower disappears--sometimes +completely, at other tunes partially. The direction of the stamens and +style, and even that of the whole flower, becomes altered from the +inclined to the vertical position. In addition to these changes, which +are those most commonly met with, the number of the parts of the flower +is sometimes augmented, and a tendency to pass from the verticillate to +the spiral arrangement manifested. Schlechtendal mentions some flowers +of _Tropaeolum majus_ in which the flowers were perfectly regular and +devoid of spurs[224], while in the double varieties, now commonly grown +in greenhouses, the condition of parts is precisely the same as in the +double violet before alluded to. Among the _Papilionaceae_ the Laburnum +and others have been noticed to produce occasionally a perfectly regular +flower in the centre, or at the extremity of the inflorescence, though +the peloria in this flower is usually irregular. In the Gentianaceous +genus _Halenia_, _H. heterantha_ is remarkable for the absence of spurs. +Amongst _Gesneraceae_, _Bignoniaceae_, _Scrophulariaceae_, and other +families of like structure, regular peloria is not uncommon. Fig. 120 +represents a case of this kind in _Eccremocarpus scaber_, conjoined, as +is frequently the case, with dialysis or separation of the petals.[225] +Many of the cultivated Gloxinias also show erect, regular, five +stamened flowers, but these are probably cases of irregular peloria. + +[Illustration: FIG. 120.--Regular peloria, _Eccremocarpus scaber_.] + +A solitary flower of _Pedicularis sylvatica_ was found by the Marquis of +Stafford near Dunrobin Castle in Sutherlandshire, in which the usual +ringent form of the corolla was replaced by the form called +salver-shaped. There were six stamens, four long and two short. Sir W. +Hooker and Mr. Borrer are stated to have found a similar flower in the +same locality in 1809.[226] + +The passage of ligulate to tubular corollas among _Compositae_ is not of +such common occurrence as is the converse change. I owe to Mr. Berkeley +the communication of a capitulum of a species of _Bidens_, in which +there was a transition from the form of ligulate corollas to those that +were deeply divided into three, four, or five oblong lobes. These then +were instances of regular peloria. + +[Illustration: FIG. 121.--Flower of _Cattleya marginata_. Lip replaced +by a flat petal.] + +In _Orchidaceae_ a similar change is not by any means infrequent; in a +few, indeed, a regular flower is the normal character, as in +_Dendrobium normale_, _Oncidium heteranthum_, _Thelymitra_, etc. Fig. +121, reduced from a cut in the 'Gardeners' Chronicle,' 1854, p. 804, +represents an instance of this kind in _Cattleya marginata_. + +From the same journal the following account of a case of peloria in +_Phalaenopsis Schilleriana_ is also cited as a good illustration of this +peculiar change. The terminal flower differed entirely from all the +others; instead of the peculiar labellum there were three petals all +exactly alike, and three sepals also exactly alike; the petals resembled +those of the other flowers of the spike, and the upper sepal also; but +the two lower sepals had no spots, and were not reflexed as in the +ordinary way: thus, these six parts of the flower were all in one plane, +and being close together at their edges, made almost a full round +flower; the column and pollen-glands were unaffected. Professor +Reichenbach also exhibited at the Amsterdam Botanical Congress, of 1865, +a flower of _Selenipedium caudatum_ with a flat lip. + +M. Gris[227] has placed on record some interesting cases of peloria of +this kind in _Zingiber zerumbet_; in the more complete forms the +androecium or staminal series was composed of six distinct pieces, the +three inner of which were fertile, while in the ordinary flower the +androecium is composed of two pieces, "a lip" and a fertile stamen. +"Is it not a matter of regret," says M. Gris, "to be obliged to call the +latter the normal flower?" + +Under this head may likewise be mentioned those cases in which the +normal, or at least the typical symmetry of the flower is restored by +the formation of parts usually suppressed; thus Moquin cites an abnormal +flower of _Atriplex[228] hortensis_ described by M. Fenzl as having a +true calyx within the two bracts that usually alone encircle the +stamens. Adanson, also cited by Moquin, found a specimen of _Bocconia_ +with a corolla. _Arum maculatum_ has likewise been met with provided +with a genuine perianth as in _Acorus_ and other Orontiads. The unusual +development of the sexual organs in diclinous flowers has been alluded +to under the head of heterogamy, and other cases where the symmetry of +the flower is rendered regular, by the development of parts ordinarily +suppressed, will be found in the chapters relating to deviations from +the usual number of organs. + +This change, or rather this persistence of a form that is usually +transient, is generally accompanied by some other alterations. Change of +direction, as has been already mentioned, is one of the most common of +these; separation of the petals (_Antirrhinum_, _Verbascum_, &c.), and +even their appearance in leaf-like guise, are not infrequent +(_Delphinium_, _Antirrhinum_, _Verbascum_, &c.) At other times +multiplication or increased number of the whorls of petals takes place, +often, but not always, at the expense of the sexual organs of the +flower. Perhaps even more frequent is the increased number of parts in +the same whorl in cases of regular peloria; thus, in the Pelargoniums +before alluded to, the parts of the flower are frequently regulated by +the number six instead of five. + +This form of peloria is most generally met with in flowers that are +placed at the end or in the centre of the inflorescence, or in such +flowers as occur singly at the end of the flower-stalk, as in +_Tropaeolum_, _Viola_, &c. It would hence seem as if the freedom from +pressure or restriction on one side allowed the flower to develop +equally in all directions, and thus to produce regularity of form. + +It is obvious, from what has been before said, that the process of +fertilisation is in many cases interfered with and altered by the change +in the conformation or the flower. + +From overlooking the occasional existence of this form of peloria, new +genera have sometimes been formed on insufficient grounds. The genus +_Aceranthus_, for instance, consists of species of _Epimedium_ in which +the customary spurs are not formed.[229] + +The occurrence both of regular and irregular peloria on the same plant +has frequently been observed in _Linaria_. It has also been remarked +that the seedlings raised from these forms are not always constant; +thus, the late Mr. Crocker, formerly foreman in the Royal Gardens, Kew, +informed me that he fertilised some flowers of a drooping Gloxinia with +their own pollen, and that when the seedlings blossomed a large number +of them produced the erect regular flowers. + +From what has been already said it will be seen that regular peloria is +closely allied to what Morren called epanody, or a return to the normal +condition. The reversion of a monstrous form to the normal one, as, for +instance, when the fern-leaved beech reverts to the normal type, was +called by the same author epistrophy.[230] + +The following are the genera in which regular peloria has been most +often observed. It must, however, be remarked that in some of the +flowers recorded as peloric there is no indication as to which form of +peloria the case should be referred to. For other illustrations refer to +chapters on Heterogamy, Number, Irregular Peloria, &c. + + *Delphinium peregrinum! + *Nigella damascena! + *Aquilegia vulgaris! + *Viola odorata! + hirta. + Epimedium, sp. + *Pelargonium zonale! + * inquinans! + Tropaeolum majus! + *Wistaria sinensis. + Lupinus. + *Cytisus Laburnum! + Trifolium repens! + *Compositae, gen. pl.! + Lonicera Periclymenum! + Streptocarpus Rexii. + *Digitalis purpurea. + *Scrophularia aquatica. + *Pentstemon. + *Linaria vulgaris! + *Antirrhinum majus! + Verbascum nigrum! + Columnea Schiedeana. + Halenia heterantha. + Galeobdolon luteum. + Prunella vulgaris! + Salvia, sp.! + Teucrium campanulatum. + Betonica alopecuros. + Eccremocarpus scaber. + Pedicularis sylvatica. + Zingiber Zerumbet. + Phalaenopsis amabilis! + Phalaenopsis Schilleriana. + Habenaria. + *Orchis morio. + mascula. + *Dendrobium, sp. + Atriplex, sp. + Cattleya Mossiae! + marginata. + Calanthe vestita! + Oncidium, sp.! + Selenipedium caudatum. + Arum maculatum. + +In addition to the references already given, further information on this +subject may be gained from consulting the following publications. See +also Irregular Peloria. + + Giraud, 'Bot. Soc. Edinb.,' Dec. 12, 1839, _Antirrhinum_. + Dareste, 'Ann. Sc. Nat.,' ser. 2, 1842, xviii, p. 220, + _Delphinium_. C. Morren, 'Fuchsia,' p. 90, _Calceolaria_, + 'Bull. Acad. Belg.,' xx, part ii, p. 57; and E. Morren, 'Bull. + Acad. Belg.,' 2nd ser., xix. p. 224, _Gloxinia_. Richard, 'Mem. + Soc. d'hist. nat.,' ii, p. 212, tab. 3. Lindley, 'Journ. Linn. + Soc.,' iii, p. 9, _Dendrobium_. Michalet, 'Bull. Soc, Bot. + France,' vii, p. 625, _Betonica_. Gubler, 'Bull. Soc. Bot. + Fr.,' ix, 81, 'Des anomalies aberrantes et regularisantes.' + Reichenbach fil. 'De pollinis orchid. genesi ac structura,' + 1852, _Oncidium_. Clos, 'Mem. Acad. Toulouse,' vi, 1862, + _Salvia_. Caspary, 'Verhandl. Phys. OEkon. Gesell. + Koenigsberg,' 1860, i, 59, _Columnea_. Bureau, 'Bull. Soc. Bot. + Fr.,' 1861, vol. viii, p. 710, _Streptocarpus_. Darwin, + 'Variation of Animals and Plants,' ii, pp. 59 and 396. Godron, + 'Ex. Bull. Bot. Soc. Fr.,' xiv, p. 165, 'Rev. Bibl.,' + _Wistaria_. Marchand, 'Adansonia,' iv, p. 172, _Lonicera_. + Baillon, 'Adansonia,' v, p. 177, 'Sur la regularite transitoire + de quelques fleurs irreg.,' shows that during the development + of some flowers which begin and end by being irregular, there + is an intermediate state when all the parts are regular. Helye, + 'Revue Horticole,' Sept., 1868, p. 327. In this last paper, + published as this sheet is going through the press, the author + states that he has raised from seed three generations of plants + of _Antirrhinum_ with regular spur-less flowers. The original + wild plant was only partially peloric, but all the flowers + produced on its descendants were regular. + +FOOTNOTES: + +[221] "On the existence of two forms of Peloria," by M. T. Masters. +'Nat. Hist. Review,' April, 1863. + +[222] Baillon, 'Adansonia,' iv. p. 149. + +[223] Similar cases are figured in 'Hort. Eystettens. Ic. Pl. Vern.' +fol. 4, f. 1, 2. _Viola martia_ multiplici flore. + +[224] 'Linnaea,' 1837, p. 128. + +[225] M. Bureau, 'Bull. Soc. Bot. Fr.,' ix, p. 91, describes two genera +of _Bignoniaceae_ in which the flowers are _normally_ regular and six +parted. + +[226] See 'Trans. Linn. Soc.,' vol. x. p. 227. + +[227] 'Ann. Sc. Nat.,' ser. 4, 1859. tom. xi, p. 264, tab. 3. + +[228] 'El. Ter. Veg.,' p. 342. + +[229] Marchand, 'Adansonia,' vol. iv, p. 127. + +[230] 'Bull. Acad. Belg.,' xvii. p. 17. "Fuchsia," p. 169. + + + + +PART II. + +PLEIOMORPHY.[231] + + +Most irregular flowers owe their irregularity to an unequal development +of some of their organs as compared with that of others. When such +flowers become exceptionally regular they do so either because +development does not keep pace with growth, and a regular flower is thus +the result of an arrest of the former process (regular peloria), or +because the comparatively excessive development, which usually occurs in +a few parts is, in exceptional cases manifested by all, hence the flower +becomes regular from the increase in number of its irregular elements. +These latter cases, then, are due to an excess of development, hence the +application of the term pleiomorphy. It must be understood that mere +increase in the number of the organs of a flower is not included under +this head, but under that of deviations from the ordinary number of +parts. + +FOOTNOTES: + +[231] [Greek: Pleios-morphosis]. + + + + +CHAPTER I. + +IRREGULAR PELORIA. + + +The term peloria was originally given by Linne to a malformation of +_Linaria vulgaris_, with five spurs and five stamens, which was first +found in 1742 near Upsal. This was considered so marvellous a +circumstance that the term peloria, from the Greek [Greek: pelor], a +prodigy, was applied to it.[232] After a time other irregular flowers +were found in like condition, and so the term peloria became applied to +all cases wherein, on a plant habitually producing irregular flowers, +regular ones were formed. The fact that this regularity might arise from +two totally different causes was overlooked, or at least not fully +recognised, even by Moquin-Tandon himself. Where a flower retains +throughout life the same relative size in its parts that it had when +those parts first originated the result is, of course, a regular flower, +as happens in violets and other plants. This kind of peloria may for +distinction sake be called regular or congenital peloria (see chapter on +that subject); but where a flower becomes regular by the increase in +number of its irregular portions, as in the _Linaria_ already alluded +to, where not only one petal is spurred, but all five of them are +furnished with such appendages, and which are the result of an irregular +development of those organs, the peloria is evidently not congenital, +but occurs at a more or less advanced stage of development. To this +latter form of peloria it is proposed to give the distinctive epithet of +irregular. + +Peloria is either complete or incomplete; it is complete when the flower +appears perfectly symmetrical, it is incomplete when only a portion of +the flower is thus rendered regular. It is very common, for instance, to +find violets or Linarias with two or three spurs, and these intermediate +stages are very interesting, as they serve to show in what way the +irregularity is brought about. In _Antirrhinum_, _Linaria_, &c., +intermediate forms show very clearly that it is to the repetition of the +form usually assumed by the petals of the lower lip that the condition +is due. This is also obvious in peloric flowers of the _Calceolaria_. +The perfect peloria of this flower is in general erect, with five +regular sepals, a regular corolla contracted at the base and at the +apex, but distended in the centre so as to resemble a lady's sleeve, +tight at the shoulder and wrist, and puffed in the centre! + +[Illustration: FIG. 122.--Peloric flower of _Calceolaria_.] + +Morren[233] describes a form intermediate between the ordinary +slipper-shaped corolla and the perfect peloria just described, and which +he calls sigmoid peloria. This flower is intermediate in direction +between the erect peloria and the ordinary reflected flower. The tube is +curved like a swan's neck and is dilated in front into two hollow +bosses, such as we see in the lower lip of an ordinary flower; beyond +these it is contracted and is prolonged into a slender beak terminating +in two hollow teeth, between which is the narrow orifice of the +corolla. The colour at the base of the tube inside is as in the perfect +peloria; while round the summit of the tube, in both cases, the +intensity of colour is greatest on the outside. Now, in a normal flower +the deepest colour is within just opposite the orifice of the corolla; +this deep colour is also seen outside of the central and most elevated +portions of the lower lip. In the peloria the deep colour at the base of +the tube represents that which is near the orifice under ordinary +circumstances, while the outer patch of colour at the apex corresponds +to that formed on the upper surface of the lower lip. On the other hand, +in peloric flowers of _Cytisus Laburnum_, _Clitoria Ternatea_, +_Trifolium repens_, and other Papilionaceae, it is the "standard," the +form of which is repeated. In the case of peloric aconites[234] the +lateral and sometimes the inferior coloured sepals assume the hooded +form usually peculiar to the upper sepal only, the number of the petals +or nectaries being correspondingly increased. Balsams become peloric by +the augmentation in the number of spurs.[235] So when orchids are +affected with irregular peloria it is the form of the labellum that is +repeated, the accessory lips being sometimes the representatives of +stamens, which are usually suppressed in these flowers,[236] but at +other times the appearance is due simply to the fact that all three +petals assume the form usually confined to the lip, the staminal column +being unaffected, except that its direction and relative position with +reference to the other parts of the flower is different from ordinary. +This was the case in some flowers of _Phalaenopsis equestris_ sent to me +by Mr. Wentworth Buller. Fig. 123 represents a flower of _Aristolochia +caudata_ with two lips, for which I am indebted to Mr. W. H. Baxter. + +From these cases it is evident that the flowers in question become +regular by the repetition of the irregular parts. + +[Illustration: FIG. 123.--Two-lipped flower of _Aristolochia caudata_.] + +It is probable that peloria may occur in any habitually irregular +flower, and that, if more attention were directed to the subject, +illustrations might be obtained from a larger number of natural families +than can be done at present. It is, however, necessary to exercise +discrimination, and not to attribute to peloria all the cases that at +first sight appear to be so referable. Thus, Professor Dickson exhibited +at the Botanical Society of Edinburgh, December 13th, 1860, four +abnormal flowers of the common Indian cress (_Tropaeolum majus_), each +presenting a supernumerary spur. On these he remarked that "in +_Tropaeolum_ the posterior part of the receptacle between the insertion +of the petals and that of the stamens is dilated so as to form the spur +which is so characteristic in the genus. The position of the spur in a +line with the posterior sepal has led many botanists to consider it as a +process of that sepal, but the fact of its being situated within the +insertion of the petals is conclusive as to its receptacular origin. In +the flowers exhibited the supernumerary spur (as if to show its want of +connection with any sepal) was placed exactly between a lateral sepal +and one of the anterior sepals, sometimes on the one side of the flower +and sometimes on the other. These additional spurs were precisely +similar to the normal ones, except that they were a little shorter. This +abnormality, although at first sight seeming to indicate a pelorian +tendency, is no approximation to regularity, from the fact of the extra +spur being differently placed, with regard to the sepals, from the +normal one." + +Peloria of this kind, when perfect, is very often associated with other +alterations. Change of direction is one of the most common of these; the +usually drooping flower becomes erect, the stamens and style also are +changed in direction, while, not unfrequently, either the one or the +other (most often the stamens) are entirely suppressed. With this +suppression an increase in the size of the flower very generally +coincides. The number of parts is also frequently increased; thus, in +_Antirrhinum majus_ the corolla, when subjected to peloria, is very +generally six-parted, and has six stamens. Fusion of one or more flowers +is also a common accompaniment of peloria, as in _Digitalis purpurea_, +in which plant prolification often adds increased complexity to the +flower. + +It has been stated by Moquin and others that the uppermost flower of an +inflorescence is the most subject to peloria; the uppermost flower of +_Teucrium campanulatum_, for instance, is very generally regular. In +_Calceolaria_ it is the central terminal flower which is usually +peloriated; on the other hand, in _Linaria_ and _Antirrhinum_ the lower +flowers, or those on the secondary branches, are quite as often affected +as the primary ones. Cassini considered that the spur of _Linaria_ was +developed from the lower petal rather than from the upper ones, because +there is more room on the side of the flower farthest from the stem than +on the opposite side. With reference to this point, M. Godron remarks +that in habitually irregular flowers the apex of the peduncle is +oblique, and hence the flowers are bent downwards or spread +horizontally, but if the receptacle be quite flat and level then the +flower is regular. The oblique position causes some of the organs to +press on others, and hence induces abortion and suppression of some +parts and increased growth in others that are not subjected to pressure. +In a terminal peloriated flower of aconite, described by this +naturalist, the flower was removed so far from the nearest bracts that +all its parts had the chance of growing regularly. In ordinary cases M. +Godron considers that the compression of the lateral bracts is the cause +of the irregularity of the androecium and of the receptacle.[237] + +It has also been somewhat too generally stated that peloria occurs +principally on luxuriant vigorous plants. It seems quite as often to +happen in plants characterised by their deficiencies in this respect. On +this point M. de Melicoq[238] says, referring to _Linaria vulgaris_ +affected with peloria, that on the weakest plants the peloriated flower +was at the top of the stem; while in stronger plants, with more numerous +flowers and larger foliage, the peloriated flowers were principally to +be found in the centre and at the base of the inflorescence, and their +pedicels were much longer than usual. + +Linne, as has been already stated, considered these flowers to be +sterile, and only capable of multiplication by division of the root, but +Willdenow obtained seeds from the _Linaria_ which reproduced the +anomaly when sown in rich soil. Baron Melicoq obtained similar +results.[239] Mr. Darwin[240] raised sixteen seedling plants of a +peloric _Antirrhinum_, artificially fertilised by its own pollen, all of +which were as perfectly peloric as the parent plant. On the other hand, +the same observer alludes to the tendency that these peloric plants have +to revert to the usual form, as shown by the fact that when the peloric +flowers were crossed with pollen from flowers of the ordinary shape, and +_vice versa_, not one of the seedlings, in either case, bore peloric +flowers. Hence, says Mr. Darwin, there is in these flowers "a strong +latent tendency to become peloric, and there is also a still greater +tendency in all peloric plants to reacquire their normal irregular +structure." So that there are two opposed latent tendencies in the same +plant. A similar remark has been made with reference to malformations in +general by other observers. + +It would be very interesting if some competent naturalist would collect +information as to whether any variations in degree of fertility exist in +the three forms of flowers in _Linaria_, viz. the ordinary one-spurred +form, which is intermediate between the spur-less and the five-spurred +form. It must be remembered, however, that in the latter cases the +stamens are often deficient. In the _Compositae_, where there are regular +flowers in the disc and irregular ones in the ray, sexual differences, +as is well known, accompany the diversities in form. + +To Mr. Darwin the author is indebted for the communication of some +flowers of _Corydalis tuberosa_ (figs. 124, 125), provided with two +spurs of nearly equal size. To these flowers allusion is made in the +work already quoted[241] in the following terms:--"_Corydalis tuberosa_ +properly has one of its two nectaries colourless, destitute of nectar, +only half the size of the other, and therefore to a certain extent in a +rudimentary state; the pistil is curved towards the perfect nectary, and +the hood formed of the inner petals slips off the pistil and stamens in +one direction alone, so that when a bee sucks the perfect nectary the +stigma and stamens are exposed and rubbed against the insect's body. In +several closely allied genera, as in _Dielytra_, there are two perfect +nectaries; the pistil is straight, and the hood slips off on either +side, according as the bee sucks either nectary." In the flowers of +_Corydalis_, which were provided with two perfect nectaries containing +nectar, Mr. Darwin considers that there has been a redevelopment of a +partially aborted organ, accompanied by a change in the direction of the +pistil, which becomes straight, while the hood formed by the petals +slips off in either direction, "so that these flowers have acquired the +perfect structure, so well adapted for insect agency, of _Dielytra_ and +its allies." + +[Illustration: FIG. 124.--Two-spurred flowers of _Corydalis_.] + +[Illustration: FIG. 125.--Section through two-spurred flowers of +_Corydalis_, Magnified.] + +Peloria, then, is especially interesting physiologically as well as +morphologically; it is also of value in a systematic point of view, as +showing how closely the deviations from the ordinary form of one plant +represent the ordinary condition of another; thus, the peloric +Calceolarias resemble the flowers of _Fabiana_, and De Candolle,[242] +comparing the peloric flowers of _Scrophulariaceae_ with those of +_Solanaceae_, concluded that the former natural order was only an +habitual alteration from the type of the latter. Peloric flowers of +_Papilionaceae_ in this way are indistinguishable from those of +_Rosaceae_. In like manner we may trace an analogy between the normal +one-spurred _Delphinium_ and the five-spurred columbine (_Aquilegia_), +an analogy strengthened by such a case as that of the five-spurred +flower of _Delphinium elatum_ described by Godron.[243] The _Corydalis_, +before referred to, is another illustration of the same fact, the +structure being the same as in _Dielytra_, &c. + +The ordinary irregular flowers may possibly be degenerated descendants +of a more completely organized ancestor, and some of the cases of +peloria may therefore be instances of reversion; some ancient _Linaria_ +may, perhaps, have had all its petals spur-shaped, and the cases of +irregular peloria now found may be reversions to that original form. +When both regular and irregular forms of peloria occur on the same +plant, as they frequently do in _Linaria_, the one may be perhaps +considered as a reversion to a very early condition, the other to a +later state, when all the petals were irregularly formed. But before we +can assert the truth of this surmise we must have better evidence as to +what the original condition really was than we have at present. + +The proximate cause of irregular peloria has been considered to be +excess of nourishment, but evidence as to this point is very +conflicting. Willdenow states that "radices peloriae, solo sterili +plantatae, degenerant in Linariam," ('Sp. Plant.,' iii, p. 254); but this +opinion is counterbalanced by that of others, while the frequent +existence of both forms on the same plant, at the same time, seems to +negative the supposition of any direct effect from external +circumstances. + +The following are the plants in which irregular peloria has been most +often observed: + + Aconitum Napellus. + Delphinium elatum! + Corydalis tuberosa. + *Viola odorata! + hirta. + Impatiens Balsamina. + Clitoria Ternatea. + Cytisus Laburnum! + Trifolium repens! + Lupinus polyphyllus! + *Gloxinia, var. cult.! + *Linaria vulgaris! + spuria. + Elatine. + triphylla. + aeruginea. + triornithophora. + pilosa. + chalepensis. + cymbalaria! + purpurea! + decumbens. + Pelisseriana. + origanifolia. + Digitalis orientalis. + * purpurea! + Calceolaria crenatiflora. + rugosa. + * var. cult.! + Chelone barbata. + *Antirrhinum majus! + Rhinanthus crista galli. + Pedicularis sylvatica. + Pedicularis euphrasioides. + Scrophularia aquatica! + Sesamum indicum. + Lamium. + Mentha. + Sideritis. + Nepeta diffusa. + Galeopsis Ladanum. + Tetrahit. + Galeobdolon luteum. + Teucrium campanulatum! + Plectranthus fruticosus. + Cleonia lusitanica. + Dracocephalum austriacum. + Phlomis fruticosa! + Vitex incisa. + Aristolochia, sp.! + Ophrys aranifera! + Orchis simia. + pyramidalis! + latifolia! + morio! + papilionacea. + mascula. + latiflora. + conopsea. + Habenaria bifolia. + Corallorhiza innata. + Aceras anthropophora. + Cattleya Moasiae! + Phalaenopsis equestris! + Pogonia ophioglossoides! + +The literature of peloria is very extensive. The following are the +principal papers, not already mentioned, which relate to the subject, +arranged under the genera, placing those first which are most subject to +this anomaly (see also Regular Peloria). + + _Linaria_.--Adanson, 'Fam. Plant.,' t. i, p. 110. Jussien, + 'Gen. Plant.,' p. 120. Poiret, 'Encycl. Method, Suppl.,' t. + iii, Jaeger, 'Missbilld. der Gewachs.,' pp. 94, 97, and 313. + Cassini, 'Op. Phytol.,' t. ii, p. 331. Ratzebourg, 'Animadv. ad + pelor. spectand.,' 1825. Turpin. 'Ic. Veget.,' tab. xx, f. 16. + Curtis, 'Flor. Londin.,' i, 118. Hopkirk, 'Flora Anom.,' pl. + vii, figs. 1, 2, 3. Haller, 'Act. Helvet.,' 2, p. 25, t. iv. De + Candolle, 'Flore Franc.,' t. iii, p. 583. Sowerby, 'Engl. + Bot.,' iv, 260, ed. Syme, tab. 963. Chavannes, 'Mon. + Antirrhin.' Delavaud, 'Bull. Soc. Bot. France,' 1858, p. 689; + id., 1860, p. 175. Heufler, 'Linnaea,' xvii, tab. ii. Weber, + 'Verhandl. des Nat. Hist. Vereins. f. d. Rh. Preuss.,' 1850, + tab. i, figs. 1-8. 'Verh. Nat. Hist. Ver. Rh. Preus.,' 1849, + vol. vi, p. 290, tab. xiii.--_Antirrhinum_, Clos, 'Mem. Acad. + Toulous.,' vi, 1862. Chavannes, 'Mon. Antirrh.,' p. 62. + Fresenius, 'Mus. Senkenb.,' ii, t. iv, fig. 10. 'Bot. Soc. + Edinb.,' 1851, July 10.--_Calceolaria_, Chamisso, 'Linnaea,' t. + vii, p. 206. Guillemin, 'Archiv. Bot.,' t. ii, p. 1 et 136. + Schlechtendal, 'Linnaea,' xii, p. 686. Ernst Meyer, 'Linnaea,' + xvi, 26, tab. iii. Morren, 'Bull. Acad. Belg.,' t. xv, n. 7, et + t. xviii, p. 583. 'Gard. Chron.,' 1850, p. 389; ibid., 1866, p. + 612.--_Viola_, Leers, 'Flor. Herborn.,' p. 145. De Candolle, + 'Organ. Veget.,' t. i, p. 519, pl. xlv. Forbes, 'Proc. Linn. + Soc.,' June 6, 1848, p. 382. Hildebrand, 'Bot. Zeit.,' 1862, + vol. xx, tab. viii.--_Orchidaceae_, His, 'Jourl. Phys.,' 65, p. + 241. Wydler, 'Arch. Bot.,' t. ii, p. 310, tab. xvi. R. Brown, + 'Obs. organ. Orchid.,' p. 698. A. Richard, 'Mem. soc. d'hist. + nat.,' t. i, p. 212. Greville, 'Flora Edinens.,' p. 87 + (_Corallorhiza_). Curtis, 'Flora Londinensis,' t. lxxxii. + Morren, C., 'Bull. Acad. Roy. Belg.,' t. xix, part ii, p. 171. + Clos, 'Mem. Acad. Sc. Toulous.,' 5 ser., vol. iii. Caspary, + 'Schrift. K. Gesellsch. Koenigsberg,' 1860, i, 59. Masters, + 'Jourl. Linn. Soc.,' vol. viii, p. 208 (_Ophrys_, _Pogonia_). + Duchartre, 'Bull. Soc. Bot. Fr.,' vol. vii, 1860, p. 26, + _Cattleya_. Cramer, 'Bildungsabweich.'--_Limosella_, Baillon, + 'Adansonia,' i, p. 305. (Flower normally irregular, becoming + regular "a force d'irregularite.")--_Chelone_, Chamisso, + 'Linnaea,' vii, p. 206,--_Clitoria_, Bonavia, 'Gard. Chron.,' + 1868, p. 1013. In this latter communication, published as this + sheet is passing through the press, the author gives an + interesting account of the transitional stages between the + ordinary papilionaceous condition and the regular form which is + like that of a Rosaceous plant. The peloric form is stated to + be transmitted by seed. + + For other references see Moq.-Tandon, 'El. Terat. Veget.,' p. + 186. Hallier, 'Phytopathol.,' p. 151. + +FOOTNOTES: + +[232] 'Amoen. Acad.,' i, p. 55, t. iii (1744):--The following note +refers to Linne's notion that these forms were due to hybridization. It +is extracted from Gmelin's edition of the 'Systema Naturae,' 1791, p. +931. "_Linariae_ proles hybrida, ejusdemque qualitatis et constans, +radicibus infinite sese multiplicans charactere fructificationis +diversissima, corolla regulari, quinque-corniculata, pentandra, ut genus +proprium absolute constitueret et distinctissimum, nisi fructus +frequentissime abortiret. Naturae prodigium. Ita quidem a Linne. +Verisimilor autem videtur ea opinio, quae peloriam pro peculiari +degeneratione monstrosa floris habet, in quam inclinare hoc genus +(Linaria) prae aliis, similis a forma deflexio in aliis speciebus, e.g. +_spurio Elatine_, _cymbalaria_, observata, ... Merk., 'Goett. gel. +Anz.,' 1774, n. 121. Linck, 'Annal. Naturg.,' i, p. 32." + +[233] 'Bull. Acad. Belg.,' xviii, part i, p. 591. Lobelia, p. 137. + +[234] See also Seringe, 'Esquisse d'une Monogr. du genre _Aconitum_,' p. +124. + +[235] Schlotterbec, 'Act. Helvet.,' t. ii, pl. i, Roeper. Balsam, p. 10, +note. + +[236] Masters. "Peloria, &c., _Ophrys aranifera_," 'Journ. Linn. Soc.,' +viii, p. 207. + +[237] Godron, "Mem. sur les Fumariees a fl. irreg.," 'Ann. Sc. Nat.,' +ser. 5, vol. ii, tab. xvii, p. 280. + +[238] 'Bull. Soc. Bot. France,' vol. v, 1858, p. 701. + +[239] 'Bull. Soc. Bot. France,' vol. vi, 1859, p. 717. + +[240] 'Variation of Anim. and Plants,' ii, p. 70. + +[241] Loc. cit., p. 59. + +[242] 'Theor. Elem.,' ed. 2, p. 266. + +[243] Cited in 'Bull. Soc. Bot. France,' vol. xiii (Rev. Bibl.), p. 81. + + + + +PART III. + +METAMORPHY. + + +Much of the objection with which Goethe's famous essay on the +'Metamorphosis of Plants' was met on its publication may be traced to a +misapprehension of the sense in which Goethe employed the word. As used +by him, it had nearly the same signification as now applied to the word +development by organogenists. It does not necessarily imply that there +has been a change in any particular organ, but rather that there has +been, to some extent, a change in the plan of construction, in +accordance with which a deviation from the customary form results. The +particular organ was never anything else than what it is; it has not +been metamorphosed in the ordinary sense of the word; for instance, in a +double flower, where the stamens are, as it is said, changed or +metamorphosed into petals, no absolute change really has taken +place--the petal was never a stamen, although it occupies the position +of the latter, and may be considered a substitute for it. + +The term metamorphosis, then, really implies an alteration in the +organizing force, taking effect at a very early period of the life of +the flower, at or before the period when the primitive aggregation of +cells, of which it is at that time composed, becomes separated or +"differentiated" into the several parts of the flower. In other words, +the "development" of the flower pursues a different course from what is +usual. In the preceding sections the effects of arrest and of excess in +this process have been partly treated of; other deviations arising from +similar causes will be mentioned elsewhere, but, under the present +heading, are specially included cases not of merely diminished or +increased, but of perverted development; the natural process is here not +necessarily checked or enhanced, but it is changed. Hence, in the +present work, the term metamorphy is employed to distinguish cases where +the ordinary course of development has been perverted or changed. As it +is applied solely for teratological purposes, the ordinary acceptation +of the term, as nearly synonymous with "development," is not interfered +with. + +In order to avoid other possible misapprehensions, the terms retrograde +and progressive metamorphosis employed by Goethe are not herein used, +their place being, to a great extent, supplied by the more intelligible +expressions arrest or excess of development.[244] + +FOOTNOTES: + +[244] See Goethe, 'Versuch. der Metam. der Pflanzen,' 1790. English +translation by Emily M. Cox, in Seemann's 'Journal of Botany,' vol. i, +1863, p. 327. For a brief sketch of the origin and progress of the +theory of vegetable morphology, prior to the publications of Wolff, +Linne, and Goethe, as well as for an attempt to show what share each of +these authors had in the establishment of the doctrine, the reader is +referred to an article in the 'Brit. and For. Medico-Chirurgical +Review,' January, 1862, entitled "Vegetable Morphology: its History and +Present Condition," by Maxwell T. Masters. + + + + +CHAPTER I. + +PHYLLODY. + + +This condition, wherein true leaves are substituted for some other +organs,[245] must be distinguished from Virescence, q. v., in which the +parts affected have simply the green colour of leaves, without their +form or structure. The appearance of perfect leaves, in place of other +organs, is frequently looked on as due to retrograde metamorphosis, or +to an arrest of development. But this is not strictly correct; for +instance, suppose a petal, which is very generally merely the sheath of +a leaf, with the addition of colouring matter, to be replaced by a +perfect leaf, one in which all three constituent parts, sheath, stalk, +and blade, are present, it surely can hardly be said that there has been +any retrogression or arrest of development in the formation of a +complete in place of an incomplete organ. The term retrograde here is +used in a purely theoretical sense, and cannot be held to imply any +actual degradation. Morphologically, as has been stated, the case is one +of advance rather than the reverse, and hence the assignment of +instances of this nature to a perversion of development, rather than to +a diminution or to an exaltation of that process, seems most consistent +with truth. The affected organs have really undergone no actual change, +simply the direction of the organising force has been altered at a very +early state, so that the usual differentiation of parts has not taken +place. + +[Illustration: FIG. 126.--'Rose plantain,' _Plantago media var._, spike +contracted; bracts leafy.] + +=Phyllody of the bracts.=--As bracts are very generally imperfect +organs, so their replacement by perfect leaves is not attributable to +arrest of development or retrograde metamorphosis, but the reverse. The +bracts of some species of _Plantago_[246] are very subject to this +change. Thus, in the rose plantain of gardens, _P. media_ (fig. 126), +the bracts are leafy and the axis depressed or not elongated, so that it +is surmounted by a rosette of small leafy organs. A similar condition of +the bracts, unattended with arrest of growth in the axis, is common in +_P. major_ (fig. 127) and in _P. lanceolata_ (see p. 108). It also +occurs in the bracts of _Corydalis solida_, _Amorpha fruticosa_, _Ajuga +reptans_, _Parthenium inodorum_, _Centaurea Jacea_, in the involucral +bracts of the dandelion, the daisy, and many other composites. In the +'Gardeners Chronicle,' 1852, p. 579, is figured a dahlia in which the +bracts of the involucre and the scales of the receptacle had all assumed +the form, texture, and venation of leaves.[247] + +[Illustration: FIG. 127.--Leaf-like bracts in _Plantago major_.] + +[Illustration: FIG. 128.--Dahlia. Scales of receptacle leafy.] + +In _Umbelliferae_ the substitution of leaves for involucral bracts is not +infrequent. It has been observed among other plants in _Angelica +Razoulzii_, _Carum carui_, _Daucus Carota_, &c. The scales of the hop +(_Humulus Lupulus_) not infrequently manifest this change, as do also +the bracts of many amentaceous plants, _e.g._ in the male catkins of +the walnut, the female catkins of the alder,[248] of some willows,[249] +&c. The bracts of some _Euphorbiaceae_, as _E. pusilla_, _E. Lathyris_, +_E. Cyparissias_, have been observed to undergo a similar +alteration.[250] + +Amongst monocotyledons an analogous change occurs not unfrequently, as +in some commelynaceous plants, _e.g._ _Tradescantia_, in _Musa_, &c. + +The spathe of _Arum maculatum_ is sometimes represented by a stalked +leaf similar to that which occurs, under ordinary circumstances, in +_Spathiphyllum_, but in which genus the spadix is more or less adherent +to the leaf-like spathe.[251] In _Schoenus cephalotes_ a similar +exaggerated development of the bracts is figured by Rottboell.[252] + +=Phyllody in inflorescence of Conifers.=--This demands passing notice by +reason of the interest attaching to the morphological construction of +these plants. The elongation of the axis which occurs in the female +cones has been already alluded to under the head of prolification of the +inflorescence. This change is frequently associated with a more or less +foliaceous condition of the bracts, which, indeed, may be seen to be +serially continuous, both above and below, with the ordinary leaves. The +scales, too, become notched and bipartite, and show, between the lobes, +the rudiment of a bud, which in a further stage becomes developed into a +shoot bearing leaves. Such a change has been described by Parlatore in +_Abies Brunoniana_, and examples may frequently be met with in the larch +(_Larix europaea_), and specially in _Cryptomeria japonica_.[253] The +scales of the male catkins of conifers likewise occasionally assume the +appearance of leaves; this may be seen in monstrous catkins of +_Araucaria_, as also in _Podocarpeae_ and _Cupressineae_ (Eichler). + +=Phyllody of the calyx.=--Sepals under ordinary circumstances are so +like leaves, that it is not wonderful that they are often replaced by +those organs.[254] A singular instance of this has been mentioned as +occurring in _Cakile maritima_, wherein the sepals were found by M. +Fournier to be pinnatifid like the ordinary leaves of the plant.[255] +The sepals of _Ranunculaceae_ and _Rosaceae_, for example, _Rosa_, _Geum_, +are particularly liable to this change. + +[Illustration: FIG. 129.--Flower of rose, sepals replaced by five +perfect leaves; axis prolonged through the flower in the form of a leafy +branch.] + +In a species of _Geranium_ recently examined the sepals presented +themselves in the form of three-lobed leaflets; so in fuchsias and in +_Epilobium hirsutum_ the sepals occasionally are not distinguishable +from ordinary leaves (fig. 130). In roses, the change in question is a +very frequent accompaniment of prolification (fig. 129). In the peach +also this replacement of the sepals is sometimes carried to such an +extent, that five perfect, bistipulate leaves occur in the place of the +calyx, but when this is the case it usually happens that the pistil is +abortive. + +[Illustration: FIG. 130.--Fuchsia, with one of the sepals leaf-like.] + +De Candolle[256] figures a curious instance wherein the pappus of +_Podospermum laciniatum_ was replaced by five linear, foliaceous lobes. +A similar change has been noticed in other composites, as in _Tragopogon +pratense_. Engelmann mentions as subject to this hypertrophy of the +pappus, as it may be termed, _Scorzonera octangularis_ and _Senecio +vulgaris_. Wigand has observed a similar transformation in a species of +_Centranthus_ (_Valerianaceae_). + +In some cases the phyllody of the sepals has a special interest, as +bearing on the question whether what is termed calyx-tube is or is not a +portion of the calyx, and whether the sepals are modifications of the +blade or of the sheath of the leaf. Thus in the primrose the phyllodic +sepals seem to show clearly that the sepals are in that plant of a +laminar nature (fig. 131). The so-called calyx-tube of roses is +elsewhere alluded to. The leaf-like organs sometimes seen at the apex of +a cucumber would seem to support the view that there was really a +calyx-tube in _Cucurbitaceae_ adherent to the carpels. It is also shown +in the cut, fig. 132, borrowed from the 'Gardeners' Chronicle,' 1859, +p. 654. + +[Illustration: FIG. 131.--Primrose. Calyx of foliaceous segments.] + +[Illustration: FIG. 132.--Leafy calyx of melon.] + +Under ordinary circumstances, the sepals may be considered as the +representatives of the sheath of the leaf (cataphyllary) or of the blade +(euphyllary), the arrangement of the veins being different in the two +cases; thus, in the vagina or sheath, there are generally several large +veins of about equal size, either convergent towards the apex, or +divergent; on the other hand, in the blade, there is usually but one +central vein, the midrib, larger than the rest, and the smaller veins +come off at a less acute angle, and are more reticulated.[257] + +Now, when phyllomorphy occurs in sepals which ordinarily are vaginal, it +is obvious that the case is one, not merely of increased relative +growth, but also of the appearance or development of an organ habitually +suppressed; on the other hand, when phyllomorphy occurs in sepals which +usually are laminar in form and nervation, the case is one of unusual +growth or hypertrophy, and not of the development of an organ habitually +suppressed, so that the amount of change is greater in the former than +in the latter instance. + +Under normal circumstances it will be found that laminar venation is +most common in gamosepalous and vaginal venation in polysepalous +calyces. And the same holds good in cases where the calyx is abnormally +leafy. The complete leaf development shows itself more frequently among +the monosepalous plants than in the polysepalous ones, as shown even in +the subjoined list of species. This statement would be more fully +verified were it possible to state the frequency with which the +condition occurred in _individual plants_, when it would be found that +phyllody of the calyx occurs much more often in individual gamosepalous +plants than in polysepalous ones. + +Phyllody of the calyx has been most often observed in the following +plants: + + Ranunculus acris! + Delphinium Ajacis. + Caltha palustris. + Anemone Pulsatilla. + sylvestris! + nemorosa! + hortensis! + coronaria! + *Papaver orientale. + Escholtzia crocea. + Cakile maritima. + Diplotaxis tenuifolia. + Thlaspi arvense. + Cheiranthus Cheiri. + incanus. + Sinapis arvensis. + Brassica oleracea! + Peltaria alliacea. + *Sisymbrium officinale. + Caryophyllaceae,[258] sp. pl. + Geranium, sp.! + *Fuchsia, var. hort.! + Epilobium hirsutum! + Cucurbita Pepo! + *Rosa, var. hort.! + Potentilla nepalensis. + Fragaria sp. + Geum rivale. + Amygdalus communis. + Persica vulgaris. + Cerasus! + Pyrus Malus. + Daucus Carota. + Athamanta Cervaria. + *Trifolium repens! + Centranthus macrosiphon. + Tragopogon pratense. + orientale. + Scorzonera octangularis. + Hypochaeris radicata. + *Senecio vulgaris! + Podospermum laciniatum. + Cirsium arvense. + Carduus heterophyllus + tataricus. + Campanula, sp. + Convolvulus sepium. + *Primula officinalis, var. cult! + acaulis. + elatior. + Gentiana campestris. + *Petunia violacea! + Lycium europaeum. + Laurus Sassafras. + Tulipa Gesneriana. + Convallaria maialis. + Colchicum autumnale! (virescent?) + + Consult also Turpin, 'Atlas de Goethe,' t. iv, f. 12, _Lycium_. + Engelmann, 'De Anthol.,' Sec. 35, p. 31. This author figures + phyllodic sepals in _Senecio vulgaris_, tab. v, figs. 24-26; + _Campanula_, tab. iii, f. 15, 16; _Athamanta cervaria_, tab. v, + f. 14. Lindley, 'Elements of Botany,' 1847, pp. 64, 73, &c. + 'Gard. Chron.,' 1858, p. 685; 1859, p. 654, _Cucurbita_. + Petunnikoff, 'Bull. Soc. Imp. Moscow,' 1862, _Cirsium_. Braun, + 'Rejuvenescence,' Ray Society's Transl. See succeeding + paragraphs. + +=Phyllody of the corolla.=--The petals also are frequently replaced by +leaves, though in many of the recorded instances the change has been one +of colour only; these latter are strictly cases of virescence. M. +Seringe[259] speaks of a flower of _Peltaria alliacea_ in which the +calyx was petal-like, while the corolla was leafy as if there had been +transposition of the two organs, a very rare, if not unparalleled, +instance. In a flower of _Campanula Medium_, provided, as is often the +case, with a double corolla, the outer corolla was slit down on one +side, the edges of the cleft being leafy. + +[Illustration: FIG. 133.--Sepals and petals to leaves. _Geranium_.] + +The frondescent petals are very often completely disjoined, as in +_Verbascum nigrum_, and _Lonicera Periclymenum_, in which, moreover, +median prolification generally coexists. In the case of _Tropaeolum +majus_, the ordinary leaves of which are peltate and orbicular, the +petals when frondescent have not the peltate arrangement, but are +spathulate, and provided with very long, narrow stalks, so that, in some +cases, they are, more properly speaking, enlarged virescent petals than +true leaves; in other instances, however, the arrangement of the veins +is more like that of the true leaves than that of the petals. + +As might be expected, frondescence of the petals is frequently +accompanied by other changes of a similar nature in other parts of the +flower, and sometimes by the abortion of the sexual organs. Thus, in +_Actaea spicata_, as observed by Fresenius, the petals were replaced by +true petiolate, palminerved, lobed leaves, the stamens and pistils +being abortive. In _Ranunculus_ the leaves that appear in the place of +the petals have no scale at their base, and in _Tropaeolum_ the calyx (or +receptacle) is free from the usual spur. + +The absolute frequency of this occurrence seems to be greatest in those +flowers which are normally polypetalous. The petals of these flowers, as +a general rule, are more like the leaf-sheaths than the leaf-blades as +to their venation, hence it would seem that the phyllomorphic condition +in these petals is a manifestation of a greater degree of organizing +force than that which occurs in those cases where the petals are +normally present in the form of contracted blades or laminae. (See the +remarks in the preceding section.) + +Frondescence of the petals has been observed most frequently in the +following cases; some, perhaps, were cases merely of virescence, q. v.; +see also under Chloranthy, Prolification. + + Ranunculus repens! + Delphinium Ajacis. + crassicaule. + Aquilegia vulgaris. + Actaea spicata. + *Brassica oleracea! + Diplotaxis muralis. + Hesperis matronalis. + Thlaspi bursa pastoris. + Sisymbrium tenuifolium. + Turritis glabra. + Raphanus sativus. + Peltaria alliacea. + Alyssum incanum. + Erysimum Barbarea. + officinale! + cheiranthoides. + Cheiranthus Cheiri. + *Dictamnus Fraxinella! + Lychnis sylvestris. + dioica! + Alsine media. + Cerastium vulgatum! + triviale. + Reseda lutea. + Phyteuma. + Malva sylvestris. + *Tropaeolum majus! + Geranium, sp.! + Triumfetta, sp.! + Epilobium hirsutum! + OEnothera striata. + Rubus, sp. + *Rosa, var. cult.! + *Trifolium repens! + Spiraea oblongifolia. + Amygdalus communis. + *Rosa! + Cerasus vulgaris! + Persica vulgaris! + Potentilla nepalensis. + Geum rivale. + Daucus Carota! + Heracleum Sphondylium. + Torilis Anthriscus. + Echinophora maritima. + Campanula rapunculoides. + glomerata. + Phyteuma spicatum. + Calendula officinalis. + Cirsium tricephalodes. + Senecio vulgaris. + Scabiosa columbaria. + agrestis. + Lonicera xylosteum. + Periclymenum. + Gentiana Amarella. + Gilia glomeriflora. + *Symphytum officinale. + Petunia violacea! + Verbascum, sp. + Antirrhinum majus! + Stachys sylvatica. + *Anagallis phoenicea? + Primula sinensis! + Polemonium coeruleum. + + See Moquin-Tandon, 'El. Terat. Veg.,' p. 203. Engelmann, 'De + Anthol.,' Sec. 38 _et seq._; tab. ii, figs. 8-14, _Gilia_; tab. + v, 23-26, _Senecio_; tab. v, f. 1-13, _Torilis_; tab. iv, f. 3, + _Erysimum_. 'Bull. Soc. Bot. Fr.,' vol. ii, 1855, p. 479, + _Primula sinensis_. Giraud, 'Edinb. Phil. Magazine,' 1839, + _Antirrhinum_. Jaeger, 'Act. Acad. Caes. Nat. Cur.,' vol. xiii, + 2, p. 1, tab. xli, _Tropaeolum_. Bischoff, 'Lehrbuch,' 11, 2, p. + 27, _note_, _Tropaeolum_. Fresenius, 'Mus. Senkenb.,' ii, 35, + tab. 4, fig. 5, _Actaea_. See also succeeding paragraphs and + sections in Chloranthy, Virescence, &c. + +=Phyllody of the stamens= happens less frequently than the corresponding +condition in the neighbouring organs. The structure of the anther is so +much removed from that of the leaf, that the change of the stamen from +its ordinary condition to that of a leaf must be regarded as indicating +a greater degree of perverted development than that which occurs in +those cases where less highly differentiated organs, such as the sepals, +petals, and pistils, are thus altered.[260] + +In all cases it is desirable to ascertain, if possible, what parts of +the stamen are thus transformed. In some Petunias the filaments are +unchanged, but in place of the anther is a small lamina, representing +precisely the blade of an ordinary leaf. Sometimes the connective only +is replaced by a leaf. One of the most interesting cases of this kind +that has fallen under the writer's observation was in _Euphorbia +geniculata_, in which, in addition to other changes mentioned under +prolification of the inflorescence, some of the stamens were partly +frondescent, half the anther being perfect, the other half leaf-like. +Another filament bore just above the usual joint three leaflets, two +lateral ones, somewhat conduplicate, and a third central one, half +anther, half leaflet. + +[Illustration: FIG. 134.--Flower of a _Petunia_, opened to show the +stamens partially replaced by stalked leaves.] + +In the case of frondescent flowers of _Tropaeolum majus_ the stamens are +usually absent or atrophied, but in other instances the filament is +present as usual, representing the stalk of the leaf, and surmounted by +a small lamina, but this latter, in place of being nearly flat, is +pinched up in the centre from back to front, and surmounted by a +two-lobed anther, so that the general appearance of the whole structure +is that of a central anther, supported at the base on each side by two +concave leaf-lobes, or it might be compared with a three-lobed leaf, the +terminal lobe represented by the anther. + +In _Jatropha Pohliana_, Muell. (_Adenorophium luxurians_, Pohl.), a +singular condition has been observed by M. Mueller (Argov.). In this +flower the anther, in place of being represented by the flat blade of a +single leaf, had the appearance as if two such blades were present and +coherent one with the other by their midribs, along their upper or inner +surfaces, which were directed towards the centre of the flower (fig. +136), thus resembling the cases of adhesion of leaves by their surfaces +already referred to (p. 33). In other cases, in the same plant, the +anther appeared as if formed by two collateral leaves, the faces looking +towards the circumference of the flower, and their margins so folded +together as to represent an open anther lobe (fig. 135). These cases are +apparently due, not to the formation and adhesion of two leaves, but +rather to the exuberant development of one leaf into two blades.[261] +The bearings of these and other similar malformations on the morphology +of the anther are alluded to under the head of petalody of the anther. + +[Illustration: FIG. 135.--Phylloid anther of _Jatropha_, after Mueller +(Arg.).] + +[Illustration: FIG. 136.--Leaf-like anther of _Jatropha Pohliana_, after +Mueller.] + +Phyllody of the stamens has been most often observed in the following +plants: + + Anemone nemorosa. + coronaria. + Delphinium crassicaule. + Nymphaea dentata. + Tropaeolum majus! + Dictamnus albus. + *Trifolium repens! + Torilis anthriscus. + Heracleum Sphondylium. + Daucus Carota + Epilobium hirsutum! + *Rosa, var. cult.! + Lonicera Periclymenum. + Anagallis arvensis. + Primula sinensis! + Petunia, var. cult. + Jatropha Pohliana. + Euphorbia goniculata. + +In addition to the foregoing there are very numerous instances of +similar substitution in chloranthic flowers. In the above list only +those cases are given wherein the leafy change is confined to the +stamens, or, at least, to a few only of the other parts of the flower. + +=Phyllody of the pistils.=[262]--This is of more common occurrence than +is the corresponding change in the case of the stamens. It is of +interest, as it sometimes serves to illustrate the morphological nature +of the pistil. Of this the double-flowering cherry is a well-known +illustration, the pistil being here represented by two small foliar +laminae, whose midribs are prolonged with a short style, terminated by an +imperfect stigma. It is usually the basal portion of the pistil, the +ovary, which is thus specially affected, the margins being also often +disunited so as to expose the ovules. These latter organs may be absent +or they may themselves be the subjects of foliaceous development. +Moquin[263] relates having found in the neighbourhood of Montpellier a +flower of a tulip the ovary of which was represented by true leaves, +which bore on their margins the ovules, and thus presented a striking +analogy with the carpels of those Sterculias, like _S. platanifolia_, +which are foliaceous in texture and open very early in the course of +their development. A similar occurrence has also been frequently +noticed in the Columbine and also in _Cruciferae_ and _Umbelliferae_. M. +Germain de St. Pierre mentions an instance wherein the carpels of _Salix +Babylonica_ were converted into two leaves, provided with stipules. All +the flowers of the catkins were similarly changed, so that it became +permanent, and resembled a branch. + +[Illustration: FIG. 137.--Rose, in which the axial portion of the flower +was elongated and the carpels were more or less replaced by leaves.] + +Substitutions of this kind form the green "eyes" or centres of certain +varieties of _Ranunculus_ and _Anemone_. + +In proliferous roses, or in cases where the central axis of the flower +is prolonged, it frequently happens that the pistils are more or less +replaced by leaves. Fig. 137, from a specimen of Dr. Bell Salter's, +given in the 'Gardeners' Chronicle,' shows the passage, from below +upwards, of the ordinary carpels to perfect leaves; the so-called +calyx-tube being completely deficient and the ovaries entirely superior. +Like most similar specimens, this one bears out the notion that what is +called the calyx-tube in roses is really an expansion and dilatation of +the top of the flower-stalk. + +[Illustration: FIG. 138.--Cucumber with leaf attached.] + +Fig. 138, for which I am indebted to Mr. S. J. Salter, represents a very +singular conformation in the cucumber, described by that gentleman in +'Henfrey's Botanical Gazette,' i, p. 208, and considered by him to be +due to the foliaceous condition of one of the three carpels of which the +fruit is composed. The portion near the peduncle was binary, while the +distal extremity of the fruit was ternary. The main difficulties +attending the acceptance of this explanation reside in the peculiar +reversed position of the leaf, and in the fact that the fruit of the +_Cucurbitaceae_ is probably of axial nature, the dilated and succulent +end of the peduncle adhering to and usually concealing the carpels; in +some cases, however, these latter project beyond the axial portion, +leaving no doubt as to the true nature of the structure in these +particular instances. + +Admitting the axial nature of the fruit, it might be supposed that in +Mr. Salter's cucumber an adventitious leaf had been given off from the +axis, but even on that supposition the reversed position offers a +difficulty, and there still remains to be explained the fact that the +proximal part of the fruit was binary in its constitution, the distal +end ternary. + +M. Norman[264] mentions a case wherein the carpels of _Anchusa +ochroleuca_ were replaced by two leaves; from this he draws the +inference that the pistil of borages and labiates is really composed of +two leaves, placed fore and aft, the margins of the leaves being +congenitally fused. This tallies well with the account given of the +development of these plants by Payer, Germain de St. Pierre, and others. + +In an Indian species of _Triumfetta_, not only were the petals +virescent, but the ovary also was much enlarged, and in some flowers it +was divided half way down into five lanceolate leaves (fig. 139), the +sepals and stamens being in their normal condition. + +In the preceding instances the foliaceous condition has pervaded the +entire pistil, or at any rate the basal portion or ovary, and it may be +noticed that the ovary is thus shown to consist in some cases of the +sheath of the leaf, as in _Aquilegia_; in other cases of the blade, as +in _Cerasus_, _Daucus_, &c. + +[Illustration: FIG. 139.--Flower of _Triumfetta_, sp., carpels +represented by five leaves.] + +There are cases, however, in which a part only of the pistillary +structure thus becomes foliaceous. Linnaeus, 'Prolepsis,' Sec. 9, +mentions some flowers of _Carduus heterophyllus_ and _C. tataricus_ in +which the style had grown into two green leaflets, and in which the calyx +and corolla were also leaf-like. A very singular instance is recorded by +Baillon,[265] wherein the pistil of _Trifolium repens_ consisted of +three carpels, either separate, or combined so as to form a one-celled +ovary with three parietal, pluri-ovulate placentae; the ovary in these +flowers was formed of the basal vaginiform part of the leaf; the three +styles were formed by the petioles, while the stigmas were represented +by trifoliolate leaves. The back of the leaf in these cases is usually +directed away from the centre of the flower. When this change occurs it +is commonly attended by an increased number of parts, as in the trefoil +just mentioned, or in the double cherry, where usually two foliaceous +carpels may be met with, and sometimes more. + +The change is also of interest when it affects such orders as the +_Umbelliferae_, which have their ovaries inferior under ordinary +circumstances; but when these organs assume a leafy condition they +become superior also, _i.e._ they are detached from the calyx. + +As regards the position of the ovules in these foliaceous pistils, they +may be placed, as in _Aquilegia_, _Delphinium_, &c., on the edges of the +carpel or on the surface, as in some flowers of _Ranunculus repens_ and +_R. Ficaria_. A similar position of the ovules is recorded in the case +of the vine (_Vitis_), where the pistil consisted of leaves bearing the +ovules on their inner surface.[266] The supposed causes of this and +other similar malformations are alluded to under the head of chloranthy, +but it may be here remarked that semi-double flowers, fertilised by the +pollen of similar flowers, are said to produce flowers with a centre of +small green leaves, this central tuft resulting from the expansion and +frondescence of the pistils. + +As this condition rarely occurs without corresponding changes in other +parts of the flower, further remarks on this subject will be found in +the chapter relating to Chloranthy. + +Phyllody of the pistil has been most frequently recorded in the +following plants: + + Paeonia officinalis. + Ranunculus repens! + *Aquilegia vulgaris! + Delphinium elatum. + crassicaule. + Ajacis. + amaenum. + Nymphaea dentata. + Sinapis arvensis! + Diplotaxis tenuifolia. + *Brassica oleracea! + *Sisymbrium officinale! + Dianthus. sp + Reseda Phyteuma. + Triumfetta, sp.! + Lychnis dioica. + Cerastium, sp.! + *Dictamnus Fraxinella! + Cerasus avium. + vulgaris! + *Rosa, var. cult.! + *Daucus Carota! + Heracleum, sp. + Epilobium hirsutum! + Lathyrus latifolius. + *Trifolium repens! + hybridum. + Melilotus, sp. + Medicago, sp. + Lonicera Periclymenum. + Carduus heterophyllus. + tataricus. + Scrophularia aquatica. + Symphytum officinale. + Anchusa ochroleuca. + paniculata. + *Primula sinensis! + Salix babylonica. + Hyacinthus, sp. + Tulipa, sp. + +Some of the above are probably cases of mere virescence rather than of +phyllody. For further illustrations, references to authorities, &c., see +under Chloranthy, Virescence, Prolification, &c. + +=Phyllody of the ovules.=--Pending the settlement of the existing +differences of opinion with reference to the morphological nature of the +ovule and its component parts, much interest attaches to the +malformations to which they are occasionally subject. Considered purely +in a teratological point of view, it seems clear that the ovular coats +are usually, if not always, of foliar nature, while the central nucleus +is an axial organ; but if this be so there still remains the question +whether the leafy coats of the ovule are processes of the carpel itself, +or distinct independent formations, like the scales of a leaf-bud; as to +this latter point, the evidence is at present very conflicting. Prof. +Al. Braun, who has devoted much attention to the subject, describes and +figures ovules of _Nigella_ and _Adonis_, wherein the outer coat of the +ovule was converted into a leafy, lobed mass, like the ordinary leaves, +and these he considers to be a portion, not of the carpel, but of the +ovular bud; he, however, hesitates to pronounce an opinion on the nature +of the pedicel of the ovule. In _Primulaceae_, wherein ovular changes are +very common, the leafy coat of the ovule would seem, from the nature of +the placenta, to be independent of the carpel. Morren, who studied the +changes in the ovules of _Primula sinensis_, applied the term +lepyrophylly ([Greek: lepyron], a scale) to the foliaceous condition of +the testa in this plant. Unger[267] describes a series of malformations +in _Primula sinensis_, consisting chiefly of reversions of the part of +the flower to leaves. The carpels were entirely absent in this case, and +the place of the free central placenta was occupied by a circle of +leaves, sometimes bearing imperfect ovules on their edges. An instance +of a similar kind has been described by A. de Candolle.[268] + +In these flowers the placenta seemed to be composed of several funiculi +soldered together, and bearing imperfect ovules. In other cases no +traces of ovules are visible, but the funiculi are in a foliaceous +condition. Moquin also alludes to a case of the same nature in _Cortusa +Mathioli_, in which the funiculi bore little rounded leaves. Brongniart +has described some malformations of _Primula sinensis_ in which the +ovules were transformed wholly or partially into small leaves with three +to five lobes.[269] Dr. Marchand[270] mentions similar changes in +_Anagallis arvensis_ and _Lonicera Periclymenum_. + +Cramer[271] figures ovules of _Primula sinensis_ in the form of stalked +leaves, often becoming infolded at the margins, and giving origin to a +small nucleus on their inner surface. + +M. Tassi[272] records an instance in _Symphytum officinale_ wherein the +ovules were replaced by two small linear leaves arising entirely from +the axis, and not from the carpels. + +In most of the foregoing illustrations the foliar portion of the ovule +must have been independent of the carpel; this independence is less +manifest, though probably as real in the cases now to be mentioned. In +_Sinapis_ and in _Brassica oleracea_ foliaceous ovules may occasionally +be seen, attached to the placenta by long stalks. No trace of the +nucleus is visible in these specimens. + +[Illustration: FIG. 140.--_Sinapis_, replum and ovules; the dotted line +shows the position of the carpels.] + +Griffith, in alluding to a similar case in _Sinapis_,[273] describes the +ovules as foliaceous, and having their backs turned away from the axis, +the raphe being next to the axis and representing the midrib the funicle +corresponding to the petiole. The outer tegument of the ovule, according +to Griffith, is a leaf united along its margins, but always more or less +open at its apex. No inversion can, therefore, really take place in +anatropous ovules, but the blade of the leaf is bent back on the +funicle, with which its margins also cohere. + +Caspary, in an elaborate paper on phyllomorphy occurring in _Trifolium +repens_, figures foliaceous ovules springing from the edge of an open, +leafy carpel. The nucleus of the ovule, in these cases, appears to +originate as a little bud from the surface of the leafy ovule (figs. +141, 142). + +[Illustration: FIG. 141.--Leafy ovules, &c., _Trifolium repens_.] + +In a species of _Triumfetta_ (see p. 260), of which I examined dried +specimens, the ovary was open and partly foliaceous; it bore on its +infolded margins ten erect leaflets, representing so many ovules; each +leaflet was conduplicate, the back being turned towards the placenta. + +[Illustration: FIG. 142.--Leafy ovules of _Trifolium repens_, showing +formation of nucleus, &c. After Caspary.] + +On the other hand, there are cases in which the leafy coat of the ovule, +in place of being a distinct organ, seems to originate from the margin +of the carpellary leaf itself--to be, as it were, a lobule or small +process of the carpel, and not an absolutely new growth. Thus, +Planchon[274], from an examination of some monstrous flowers of _Drosera +intermedia_, was led to the inference that the ovules are analogous to +hairs on the margins of the leaves. This acute botanist was enabled to +trace all the gradations between the simple cup formed by the +confluence of four glanduliferous hairs and the concave leaf and the +perfect ovule. + +Brongniart[275] records ovules of _Delphinium elatum_ existing in the +form of marginal lobes of the carpellary leaf itself; so that each ovule +corresponds to a lobe or large tooth of this leaf, the funiculus, as +well as the raphe, being formed by the median nerve of the lateral lobe. +M. Clos[276] mentions a similar instance in _Aquilegia Skinneri_; and +another is figured in Lindley's 'Elements of Botany,' p. 88, f. 180. + +[Illustration: FIG. 143.--Portion of an open foliaceous carpel of +_Delphinium_, with ovules on the lobules.] + +Cramer[277], from an examination of several ovular malformations, as +well as from the investigation of the mode of evolution of the ovules, +is led to a similar conclusion with reference to the production of +ovules from the modified lobes of the carpellary leaf. Figs. 143-145, +copied from Cramer, show how the nucleus of the ovule is formed as a new +growth from the surface of the lobes of the leaf in _Delphinium elatum_. + +[Illustration: FIG. 144.--Section through marginal lobe of carpel +(_Delphinium_), showing the nucleus (_n_).] + +[Illustration: FIG. 145.--Section through marginal lobe of carpel, +showing nucleus and tegument (_Delphinium_).] + +[Illustration: FIG. 146.--1. Placenta of _Dianthus_, bearing ovules and +carpels. 2. One of the ovaries separated.] + +[Illustration: FIG. 147.--Ovules of _Dianthus_ passing into carpels.] + +One of the most singular instances of ovular malformation in record is +that cited by the Rev. M. J. Berkeley, in the 'Gardener's Chronicle,' +September 28th, 1850, p. 612. The plant was a carnation, and its +placenta bore, not only ovules, but also carpels (fig. 146), the latter +originating in a perverted development of the former, so that many +intermediate stages could be traced between the ordinary ovule and the +ovary (fig. 147, 1, _a_, 2, _b_). Some of these carpels, thus derived +from the ovules, themselves bore secondary ovules on a marginal +placenta, as shown in the sections at _c_, _d_, _e_. Could such a change +occur in the animal kingdom, there would be the unfertilised ovum +converted into an ovary, and this again bearing Graafian vesicles! In +Mr. Berkeley's carnation the change was not so great, seeing that the +nucleus of the ovule was not developed, and sufficient evidence has been +above given as to the foliar nature of the primine, while for a leaf to +be folded up so as to form a carpel is an ordinary occurrence. + +It is worthy of remark that in these foliaceous ovules there is never +more than one coat, the secondine and other integuments do not make +their appearance in these cases, and that very generally the change in +question accompanies a similar foliaceous condition in the carpel, the +margins of which are more or less disunited. + +Prof. A. Braun remarks that up to this date no such change has been +observed in the ovules of Monocotyledons. + +=Changes in the nucleus of the ovule.=--The preceding remarks have had +reference especially to the ovular coats, but it is desirable also to +allude to certain points connected with the nucleus. Very frequently, +when the coat of the ovule is phylloid, as before described, the nucleus +is altogether wanting, though sometimes it is present as a small +cellular papilla; very rarely is it to be found in its perfect state. +Occasionally the nucleus is present in the guise of a small elongated +branch. Wigand cites ovular buds in every stage of progress into a +branch, sometimes even bearing indications of anthers. Wydler has +observed a similar occurrence in ovules of _Alliaria officinalis_, and +Schimper has described and figured specimens of _Nigella damascena_ in +which the outer coats of the ovule were but little changed, while the +nucleus was replaced by a leafy shoot. On one of the leaves of this +latter was found an imperfect ovule--an ovule on an ovule! + +Fig. 148 shows a floret of a species of _Gaillardia_, in which the ovule +was replaced by a leafy shoot which had made its way through a chink in +the ovary. In this specimen, however, there was no evidence to show +whether the shoot in question was a perverted development of the +nucleus, or whether it was wholly independent of the ovule. + +[Illustration: FIG. 148.--Floret of _Gaillardia_, showing leafy shoot +occupying the place of the ovule.] + +From this occasional elongation of the nucleus, as well as from the +foliar nature of the ovular coats, Prof. Alex. Braun arrives at the +conclusion that the ovule is to be looked on as a bud, the ovular +coatings, so often variable in number, representing the scales of the +bud, the nucleus corresponding to the end of the axis or growing point. +Griffith had previously expressed the same opinion from his observations +on malformed ovules of _Sinapis_ and _Lonicera_, while Caspary's +conclusions from the foliaceous ovules of _Trifolum repens_ are somewhat +similar. The latter observer considers that the funiculus, with the +integuments, is the equivalent of a leaflet, the petiolule or midrib of +which answers to the funiculus, and its hollow expansion to the +integument. The nucleus itself is considered to be a new formation +analogous to a shoot. + +M. van Tieghem's conclusion[278] from the examination, of flowers of +_Tropaeolum majus_, in which the ovules were replaced by perfect peltate +leaves, is that the ovules are foliar productions springing, not +directly from a prolonged floral axis, as in _Primulaceae_, but from +branches of the axis arising from the axils of the carpellary leaves. + +Phyllody of the ovules has been met with most often in the following +species: + + *Aquilegia vulgaris! + Skinneri. + Delphinium crassicaule. + elatum. + dictyocarpum. + Ajacis. + Nigella damascena. + Adonis autumnalis. + Cheiranthus Cheiri! + Nasturtium, sp. + Sisymbrium officinale! + Brassica napus! + * olcracea! + *Alliaria officinalis! + Sinapis arvensis! + Turritis, sp. + Thlaspi arvense. + Erucastrum Pollichii. + Stellaria media. + *Reseda lutea. + Drosera intermedia. + Agrostemma Githago. + Stellaria media. + Triumfetta, sp.! + Tropaeolum majus! + Dictamnus albus. + Fraxinella! + Caram carui + Pastinaca sativa. + Torilis anthriscus. + Thysselinum palustre. + Epilobium palustre. + Rosa, sp. + Fragaria alpina. + *Trifolium repens! + Medicago maculata. + Desmodium canadense. + Melilotus macrorhiza. + Lonicera, sp. + Gaillardia! + Crepis, sp. + Phyteuma odorata. + Symphytum Zeyheri. + * officinale. + Stachys sylvatica. + Anagallia arvensis. + phoenicea. + Lysimachia ephemerum. + *Primula sinensis! + Auricula. + praenitens. + Gilia glomeruliflora. + Rumex arifolius. + scutatus. + Salix capraea. + +The following list of publications relating to ovular malformations is +copied from A. Braun, 'Ueber Polyembryonie und Keimung von Caelobogyne' +(Appendix),[279] to which are also added some others not alluded to by +that author and not specially referred to in the preceding pages: + + Jaeger, 'Missbilld. d. Gewaechse,' p. 78, 79, f. 47. Roeper, + 'Enum. Euphorb.,' 1824. p. 45, _Delphinium_.--Schimper, + 'Flora,' 1829, pp. 437-8, et 'Mag. fur Pharmacie de Geiger,' + 1829-30, pl. iv-vi, text wanting, _Primula_, _Reseda_, + _Cheiranthus_.--Engelmann, 'De Antholysi,' 1832.--Valentin, + 'Act. Acad. Nat. Cur.,' 1839, p. 225, _Lysimachia_.--Unger, + 'Act. Acad. Nat. Cur.,' xxii, 11, 1850, p. 543, t. 5 B, + _Primula_.--'Flora (B. Z.)', 1842, p. 369, t. ii, + _Trifolium_.--Brongniart, 'Ann. Sc. Nat.,' 1834, ii, p. 308; + also 'Archives Mus. d'Hist. Nat.,' 1844, t. iv, p. 43, pl. iv, + v, _Primula_.--Reissek, 'Linnaea,' xvii, 1843, + _Alliaria_.--Wydler, 'Denkshrift. d. Regensb. Bot. Gesell.,' + 1855, iv, s. 77, t. vii, _Alliaria_.--Wigand. 'Grundlegung der + Pflanzen Teratol.,' 1850, p. 39, _Turritis_.--Wigand, 'Bot. + Untersuchungen,' 1853, p. 23, _Rosa_, _Turritis_, + _Crepis_.--Germain de St. Pierre, 'L'lnstitut,' 1853, n. 1051, + p. 351.--Rossmann, "Entwicklung der Eiknospen aus dem + Fruchtblatte," &c., 'Flora,' 1855, pp. 647 and 705.--Dareste, + 'Ann. Sc. Nat.,' 1842, p. 220, _Delphinium_.--Fresenius, 'Mus. + Senkenb.,' ii, p. 39, t. iv, f. 9, _Primula_.--Schultz, 'Flora + o. d. Bot. Zeit.,' 1834, xvii, p. 121, _Nasturtium_.--Seringe + and Heyland, 'Bull. Bot.,' 1-7, _Diplotaxis_.--Clos, 'Mem. + Acad. Toulouse,' vi, 1862, _Delphinium_.--Morren, C., 'Bull. + Acad. Belg.,' xix, part ii, p. 519, _Primula_.--Caspary, + 'Schrift. d. Physik. OEk. Gesell. zu Koenigsberg,' band ii, p. + 51, tabs. ii, iii. Fleischer, 'Ueber Missbildungen + Verschiedener Cultur Pflanzen.,' &c., Esslingen, 1862. Cramer, + 'Bildungsabweich,' p. 68, &c. &c., _Trifolium._--Moquin-Tandon, + 'El. Terat. Veg.,' p. 206, _Cortusa_.--Guillard, 'Bull. Soc. + Bot. Fr.,' 1857, vol. iv, p. 761, _Stellaria_.--Moelkenboer, + 'Tijdschrift v. Natuurl. Geschied.,' 1843, p. 355, t. vi, vii, + _Primula_.--Van Tieghem, 'Bull. Soc. Bot. Fr.,' 1865, p, 411, + _Tropaeolum_. + +=Phyllody in accessory organs.=--In addition to the ordinary organs of +the plant, what are termed the accessory organs, such as hairs, spines, +&c., sometimes become foliaceous. It is not to be wondered at that +spines, when they represent the framework of a leaf, become sometimes +clothed with cellular tissue, and thus become indeed true leaves. This +happens occasionally in _Berberis;_ a similar thing occurs in the +stipules of some _Leguminosae_; the scales of some begonias; the tendrils +of _Bignonia_, _Cobaea_, &c. + +The presence of two small green laminae on the outer side of the two +posterior stamens in _Antirrhinum majus_ has also been met with. The +adventitious organs appeared as if they were developments from the +thalamus--a kind of foliaceous disc, in fact. + +[Illustration: FIG. 149.--Leafy petal of _Epilobium_.] + +[Illustration: FIG. 150.--Chloranthy, &c. _Epilobium hirsutum_.] + +=Chloranthy.=--The term phyllomorphy is applied to the individual parts +of the flower which assume the form and appearance of leaves. By +chloranthy it is to be understood that all, or the great majority of the +organs of the flower assume these conditions.[280] In chloranthy, as +here defined, there is no unusual number of buds, as there is in +prolification, but the appearance of the flower-bud is so changed as to +make it resemble more closely a leaf-bud than a flower-bud. There is not +necessarily any increase in the number, or any alteration in the +position of the buds, but the form and appearance of the latter differ +from what is usual. Chloranthy, then, is a more complete form of +frondescence. Owing to the vagueness with which the word has been +applied by various authors, it becomes very difficult to ascertain +whether the recorded instances of chloranthy were really illustrations +of what is here meant by that term, or whether they were cases of mere +virescence (green colour, without other perceptible change), or of +prolification (formation of adventitious buds). It is, therefore, quite +possible that some of the instances to be now mentioned were not +strictly cases of chloranthy. + +[Illustration: FIG. 151.--_a._ Open leafy carpel of "green rose," with +two deformed ovules. _b._ Ovule separate. _c._ Primine removed. _d._ +Secondine and nucleus, with the bulbous end that projects through the +micropyle.] + +Seringe[281] has described a malformation in _Diplotaxis tenuifolia_ in +which all the floral organs were replaced by sixteen distinct leaflets +which had preserved their proper relative position. The _Cruciferae_, of +which family the last-named plant is a member, are particularly liable +to this malformation, as also are the _Rosaceae_, as will be seen from +the following illustrations. Roses indeed often exhibit alterations of +this kind as the commencement of prolification. There is also in +cultivation a rose[282] called the green rose, "Rose bengale a fleurs +vertes," in which all the parts of the flower are represented by leaves. +One of the most remarkable features in this plant is, that the carpels +have often two ovules on their margins. Now, Payer, in his +"Organogenie," has shown that at a certain period of the development of +the ordinary rose flower the ovary contains two collateral ovules, of +which one becomes in process of time suppressed.[283] _Geum coccineum_ +has been found by Wigand with its flowers in this condition.[284] + +Lindley[285] figures a very interesting illustration in _Potentilla +nepalensis_, in which some of the flowers have their component parts +leafy, in others the receptacle lengthens, till in extreme cases the +whole of the floral apparatus is represented by a branch bearing a +rosette of leaves. + +A particular variety of the Alpine strawberry is also described as +occasionally subject to this transformation. In these flowers the calyx +remains normal, while all the other parts of the flower, even to the +coating of the ovule, assume a leaf-like condition.[286] + +Among _Leguminosae_ a partial leafy condition (frondescence), or a more +complete degree of the same change, (chloranthy) is not infrequent, +particularly in _Trifolium repens_. In this species the changes are so +common, so various and important, that they may be alluded to in some +little detail. M. Germain de Saint Pierre,[287] in commenting on the +frequency with which the flowers of this plant are more or less +frondescent, remarks that although all the flowers on one plant may be +affected, they are all changed in the same manner, but on different +specimens different degrees of transformation are found. In all the +corolla and stamens are comparatively little removed from the ordinary +form, the calyx and pistil, however, have a particular tendency to +assume a foliar condition. The author just cited arranges the +malformations of this plant under three heads, as follows: + + 1. Calyx-teeth larger than usual, sometimes dentate at the + margin; petals more or less regular and disposed to run away + from the papilionaceous form; filaments free; anthers normal; + carpel transformed into a true leaf with a long stalk provided + at the base, with two stipules, terminal leaflet, solitary, + green, with no trace of ovules. Sometimes a second carpellary + leaf, similar to the first, is formed; in other cases the + central axis of the flower is occasionally prolonged into a + head of young flowers--median prolification. In some few + instances the calyx is not at all altered, but the carpellary + leaf is trifoliolate, or even quinquefoliolate, the corolla + being then absent. The heads of flowers in this first form have + the aspect of little tufts of leaves. + + 2. Each of the teeth of the calyx is represented by a long + stalk, terminated by a single articulated leaflet, the + bi-labiate form of the calyx is still recognisable; the two + upper petals are united, the three lower separate; the tube of + the calyx is not deformed and seems to be formed of the + petioles of the sepals united by their stipules. In this second + class of cases the corolla is papilionaceous, the filaments + free, the carpellary leaf on a long stalk provided with + stipules, its blade more or less like the usual carpel, with + its margins disunited or more commonly united with the ovules + in the interior, sometimes represented by a foliaceous, dentate + primine only. In one case the carpel was closed above, gaping + below, where it gave origin to several leaflets, the lower ones + oval, dentate, like ordinary leaflets, the upper ones merely + lanceolate, leafy lobes, representing the primine reduced to a + foliaceous condition. Inflorescence--a head with leafy flowers + on long stalks, which are longer at the circumference than in + the centre. + + 3. Calyx-teeth lance-shaped, acuminate; corolla more or less + regular, arrested in its development and scarcely exceeding the + tube of the calyx within which it is crumpled up; stamens but + little changed; carpellary leaf on a short stalk, not exceeding + the calyx tube, but the ovarian portion very long, and provided + with abortive ovules. + + These three groups will be found to include most of the forms + under which frondescence of the clover blossoms occurs, but + there are, of course, intermediate forms not readily to be + grouped under either of the above heads. Such are the cases + brought under the notice of the British Association at + Birmingham in 1849 by Mr. R. Austen, in some of which the + petals and stamens even were represented by leaves. + +Although, on the whole, chloranthy is most frequent in the families +already alluded to, yet it is by no means confined to them, as the +examples now to be given amply show. Specimens of _Nymphaea Lotus_ have +been seen in which all the parts of the flower, even to the stigmas, +were leafy, while the ovules were entirely wanting. + +Planchon[288] figures and describes a flower of _Drosera intermedia_ +that had passed into a chloranthic condition, excepting the calyx, which +was unchanged; the petals, like the valves of the ovary, were provided +with stipules, and were circinate in vernation. + +M. A. Viaud-Grand-Marais[289] records an interesting example of +chloranthy, in which the sepals, petals, pistils, and ovules of +_Anagallis arvensis_ were all foliaceous. Similar changes have not +unfrequently been met with in _Dictamnus Fraxinella_. + +M. Germain de Saint Pierre has also recorded the following deviations in +the flowers of _Rumex arifolius_ and _R. scutatus_; in these specimens +the calyx was normal, the petals large, foliaceous, shaped like the +stem-leaves, the stamens were absent, the three carpels fused into a +triangular leafy pod, as long again as the perianth, the stigmas normal +or wanting, the ovule represented by a thick funicle, terminated by a +foliaceous appendage analogous to the primine.[290] + +In grasses it frequently happens that the flowers are replaced by +leaf-buds; this condition is alluded to elsewhere under the head of +viviparous grasses, but in this place may be mentioned a less degree of +change, and which seems to have been a genuine case of chloranthy in +_Glyceria fluitans_, the spikelet of which, as observed by Wigand,[291] +consisted below of the ordinary unchanged glumes, but the remaining +paleae as well as the lodicles and stamens were represented by ligulate +leaves. The plant, it is stated, was affected by a parasitic fungus. On +the other hand, General Munro, in his valuable monograph of the +_Bambusaceae_,[292] refers to an illustration in which "the lowest glumes +generally, and the lowest paleae occasionally, had the appearance of +miniature leaves, with vaginae, ligules and cilia, enveloping, however, +perfect fertile spiculae; as progress is made towards the top of the +spike, the ligule first, then the cilia, and finally, the leaf-like +extension disappears, and the uppermost glumes assume the ordinary shape +and form of those organs." + +=General remarks on chloranthy and frondescence.=--Moquin remarks with +justice that the position of the flowers on the axis is of importance +with reference to the existence of chloranthy. Terminal flowers are more +subject to it than lateral ones, and if the latter, by accident, become +terminal, they seem peculiarly liable to assume a foliaceous condition. +Kirschleger says, that in _Rubus_ there are two sorts of chloranthy, +according as the anomaly affects the ordinary flowering branches, or the +leafy shoots of the year, the summits of which, instead of developing in +the customary manner, terminate each in one vast and long inflorescence, +very loose and indeterminate, and with axillary flowers.[293] + +On the whole, taking in consideration cases of partial frondescence, as +well as those in which most of the parts of the flower are affected, +phyllody would seem to be most common in the petals and carpels, least +so in the case of the stamens and sepals. It is more common among +polysepalous and polypetalous plants than in those in which the sepals +or petals are united together. + +The causes assigned for these phenomena are chiefly those of a nature to +debilitate or injure the plant; thus it has been frequently observed to +follow the puncture of an insect. M. Guillard[294] gives an instance in +_Stellaria media_ where the condition appeared to be due to the attacks +of an insect _Thrips fasciata_. Still more commonly it arises from the +attacks of parasitic fungi, _e.g._ _Uredo candida_, in Crucifers, &c. + +In other cases it has been observed when the plants have been growing in +very damp places, or in very wet seasons, or in the shade, or where the +plant has been much trampled on. This happens frequently with _Trifolium +repens_. The frequency with which the change is encountered in this +particular species is very remarkable; it is difficult to see why one +species should be so much more subject to the kind of change than +another of nearly identical conformation. + +It might at first be supposed that the same causes that bring about the +complete substitution of leaf-buds for flower-buds (see Heterotaxy) +would operate also in the partial substitution of leaves for other parts +of the flower, but it will be seen that the inducing cause, whether +similar or not in the two cases respectively, acts at different times; +in the one case, it is not brought into play until the rudiments of the +flower are already formed, whereas in the other the influence is exerted +prior to the formation of the flower. So that while the formation of +leaf-buds in place of flower-buds may be and generally is due to an +excess of nutrition, inducing over activity of the vegetative organs, +the production of phyllomorphic or chloranthic flowers may be owing +rather to a perversion of development arising from injury or from some +debilitating agency. The discrepancies in the assigned causes for the +conditions above mentioned may, therefore, in great measure, be +attributed to the different periods at which the causes in question +operate. + +The following list may serve as a guide to the plants most frequently +the subjects of chloranthy, but reference should also be made to +preceding and subsequent sections, and to that relating to prolification +of the inflorescence. + + Aquilegia vulgaris. + Chelidonium majus. + Corydalis aurea. + Nymphaea Lotus! + *Brassica oleracea! + Bunias. + Hesperis matronalis. + *Sinapis arvensis! + Sisymbrium officinale. + Erucastrum canariense. + Diplotaxis tenuifolia. + Lychnis dioica! + Cerastium glomeratum! + triviale. + Stellaria media. + Poterium polygamum. + Torilis anthriscus. + Seseli, sp. + Selinum caruifolium. + Epilobium hirsutum! + Begonia fuchsioides. + Gomphia, sp. + Scabiosa Columbaria. + Dipsacus fullonum. + Matricaria Parthenium. + Calendula officinalis. + Campanula pyramidalis. + Reseda odorata! + Vitis vinifera. + Dictamnus Fraxinella! + Triumfetta, sp.! + *Tropaeolum majus! + Rhamnus Frangula. + *Trifolium repens! + Lupinus, sp. + Rosa diversifolia! + Potentilla nepalensis. + argentea. + Fragaria vesca! + Geum rivale. + Rubus fruticosus. + caesius. + Saxifraga foliosa. + Verbascum phlomoides. + Scrophularia nodosa. + aquatica! + *Primula sinensis! + Lysimachia Ephemerum. + Anagallis arvensis. + Webbiana. + Nicotiana rustica. + Anchusa ochroleuca. + Myosotis caespitosa. + Stachys sylvatica. + Gilia capitata. + Euphorbia segetalis. + Rumex arifolius. + scutatus. + Juncus lampocarpus. + uliginosus. + +In addition to the publications before cited the following may be named +as containing valuable information on the subject of this chapter. + + Jaeger, 'Missbild. Gewaechs.,' 1814, p. 83, _Trifolium repens_. + For other accounts of similar malformations in the same plant, + see Schmitz, 'Linnaea,' xv, p. 268. Unger, 'Flora' (B. Z.) xxv, + p. 369. Caspary, 'Schrift. der. Physik. oekon. Gesellsch. zu + Koenigsberg,' 2, 1861, p. 51, tabs. ii, iii. Fleischer, + 'Missbilld. verschied. Cult. Pflanz.,' 1862, p. 55, &c., t. v, + vii, &c. For _Primula_ see Brongniart, 'Ann. Sc. Nat.,' ser. 2, + t. i, p. 308. A. P. and Alph. De Candolle in 'Neue + Denkschrift.' Morren, C., 'Bull. Acad. Roy. Belg.,' xix, part + 2, p. 539. Molkenboer, 'Tijdschr. voor Natuurl. Geschied.,' + 1843, p. 355, tabs. vi, vii. Marchand, 'Adansonia,' iv, p. 167 + and p. 159. _Anagallis_, p. 171, _Lonicera_, p. 83, _Juncus_. + For other plants see Fresenius, 'Mus. Senk.,' 2, p. 35, &c. + Norman, 'Ann. Sc. Nat.,' ser. 4, 1858, vol. ix, p. 220. Christ, + 'Flora' (B. Z.) 1867, p. 376, tabs. v, vi, _Stachys_. Cramer, + 'Bildungsabweich.,' p. 26, &c. Baillon, 'Adansonia,' ii, p. + 300. Moquin-Tandon, 'El. Ter. Veg.,' p. 230. Schauer's + translation, p. 220. Hallier, 'Phytopathologie,' p. 160. + +FOOTNOTES: + +[245] Engelmann makes use of the word frondescence in the same cases. +'De Anthol.,' p. 32, Sec. 38, while Morren adopts the term Phyllomorphy, +'Lobelia,' p. 95. + +[246] See Schlechtendal, 'Bot. Zeit.,' vol. xv, 1857, p. 873; also +Marchand, 'Adansonia,' iv, p. 156. + +[247] For instances of similar changes in _Composites_, see De Candolle, +'Prod.,' t. vi, p. 571, _Centaurea Jacea phyllocephala_. Clos, 'Ann. Sc. +Nat.,' ser. iii, tom. xvi, 1851, p. 41. 'Science Gossip,' 1865, p. 104, +&c. + +[248] Kickx, 'Bull. Acad. Belg.,' t. xviii, part 2, p. 288. + +[249] Weber, 'Verhandl. Nat. Hist. Vereins. f. Preuss.,' &c., 1860, p. +381. + +[250] Weber, loc. cit. + +[251] Sauter, 'Flora v. Bot. Zeit.,' 1831, p. 11. + +[252] 'Descr. et Icon. Plant.' tab. 20. + +[253] For references see p. 115; see also to Eichler, 'Excurs. +Morpholog. de format. flor. Gymnosperm.,' in "Mart. Flor. Brasil," +abstracted in English in 'Natural History Review,' April, 1864. + +[254] "Calyx tunc plane non differt a foliis proxime ipsi +praecedentibus." Wolff, 'Theor. Gener.,' Sec. 114. Linn., 'Proleps.,' +Sec. 6. Goethe, 'Versuch.,' Secs. 31-38. + +[255] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 697. + +[256] 'Organ. Veget.,' t. i, p. 492, pl. xxxii, f. 6. + +[257] This distinction between laminar and vaginal venation is well seen +in cases like _Mussaenda_, _Calycophyllum_, or _Dipterocarpus_, where +the enlarged calycine segment has a strictly vaginal arrangement of its +veins, very different from that which occurs in the true leaf-blades. +These are cases, therefore, where the sheath of the leaf is unusually +enlarged, and are not to be referred, as is often done, to metamorphosis +of one or more sepals to perfect leaves. Prolified roses, cherries, &c., +furnish frequently parallel cases. With reference to _Mussaenda_, C. +Morren held the view that the petal-like sepal was really a bract +adherent to the calyx, and incorporating with itself one of the calycine +lobes--"soudee au calice et ayant devoree, en englobant dans sa propre +masse, un lobe calicinal." The Belgian _savant_ considers this somewhat +improbable explanation as supported by a case wherein there were five +calyx lobes of uniform size, and a detached feather-veined leaf +proceeding from the side of the ovary lower down ('Bull. Acad. Belg.,' +xvii, p. 17, _Fuchsia_, p. 169). + +[258] In this order _Agrostemma Githago_ offers an illustration of a +normally leafy calyx. + +[259] 'Bull. Bot.,' i, p. 6. + +[260] Wolff's original opinion was that the stamens were equivalent to +so many buds placed in the axil of the petals or sepals (see 'Theoria +Generationis,' 1759, Sec. 114)--an opinion which more recently has received +the support of Agardh and Endlicher. Wolff himself, however, seems to +have abandoned his original notion, for in his memoir, "De formatione +intestinorum praecipue tum et de amnio spurio aliisque partibus embryonis +gallinacei, nondum visis," &c., in 'Comm. Acad. Petrop.,' xii, p. 403, +anno 1766, he considers the stamens as essentially leaves. See also +Linn. 'Prolepsis,' Sec. viii; Goethe, 'Metam.,' Sec. 46. + +[261] Mueller (Argov.), in 'Mem. Soc. Phys. et d'Hist. Nat. Genev.,' t. +xvii. + +[262] "If we keep in view the observations which have now been made, we +shall not fail to recognise the leaf in all seed-vessels, +notwithstanding their manifold forms, their variable structure, and +different combinations."--(Goethe, 'Metam.,' Sec. 78.) Wolff, 'N. Comm. +Acad. Petrop.,' 1766, xii, p. 403, expresses precisely the same opinion +as to the nature of the seed-vessel. + +[263] 'El. Terat. Veg.,' p. 205. + +[264] 'Ann. Sc. Nat.,' 4th series, vol. ix, p. 209. + +[265] 'Adansonia,' iv, p. 70. A similar deviation has been observed by +M. van Tieghem in the ovary of _Tropaeolum majus_, 'Bull. Soc. Bot. Fr.,' +1865, p. 411. + +[266] Planchon et Mares, 'Ann. Sc. Nat.,' ser. 5, vol. vi, 1866, p. 228, +tab. xii. + +[267] 'Act. Acad. Nat. Cur.,' 22, 11. 1850, p. 543, t. v, vi. + +[268] 'Neue Denkschrift der allg. Schweiz. Gesellsch.,' band v. p. 9, +tab. 3, 4. + +[269] 'Ann. Sc. Nat.,' 2 ser., vol. i, p. 308, pl. ix, c. + +[270] 'Adansonia,' vol. iv, pp. 159, 171. + +[271] 'Bildungsabweichungen,' &c., tab. iv, figs. 1, 2, 21, 28, 29, &c. + +[272] 'Bull. Soc. Bot. France,' viii, p. 395. + +[273] 'Notulae,' p. 125, atlas, pl. xxxv; and 'Journals of Travels,' +1847, p. 475, _Lonicera_. + +[274] 'Ann. Science Nat.,' 3rd ser., vol. ix, p. 86, tabs. 5, 6. + +[275] 'Comptes Rendus,' vol. xviii, March 25th, 1864, and 'Ann. Sc. +Nat.,' 3 ser., vol. ii, p. 32. + +[276] 'Mem. Acad. Sc. Toulous.,' ser. 5, vol. iii. + +[277] 'Bildungsabweich. Pflanz. Famil.,' p. 89, tab. xi. + +[278] 'Bull. Soc. Bot. Fr.,' 1865, p. 411. + +[279] Translated in 'Ann. Sc. Nat.,' 4th series, t. xiv, p. 24. + +[280] The calyx is not unfrequently excepted. + +[281] 'Bull. Bot.,' t. i, p. 6. + +[282] Lindley, 'Theor. Horticult.,' ed. 2, p. 84, f. 17. + +[283] Gris, 'Bull. Soc. Bot. Fr.,' 1858, vol. v, p. 261, and 'Ann. Sc. +Nat.,' ser. 4, vol. ix, p. 80. Planchon, 'Flore des Serres,' vol. i, +1856, p. 129. + +[284] 'Flora,' 1856, p. 711. + +[285] 'Theory of Horticult.,' ed. 2, p. 90, f. 25. + +[286] As considerable interest attaches to the "Plymouth strawberry," +and very little is known of it in this country, or on the continent, the +author gladly avails himself of this opportunity of inserting an account +of it, for which he is indebted to the kindness of Dr. Robert Hogg.--The +Plymouth Strawberry (_Fragaria vesca fructu hispido_) is a sort of +botanical Dodo upon which many have written, and which few have seen. +Many years have elapsed since it was first discovered; and although a +century and a half have passed since there was any evidence of its +existence, it serves still as an illustration for students in morphology +of one of those strange abnormal structures with which the vegetable +kingdom abounds. + +It is to old John Tradescant we are indebted for the earliest record of +this plant. Johnson, in his edition of 'Gerard,' says; "Mr. John +Tradescant hath told me that he was the first that tooke notice of this +strawberry, and that in a woman's garden at Plimouth, whose daughter had +gathered and set the roots in her garden, in stead of the common +strawberry; but she, finding the fruit not to answer her expectation, +intended to throw it away; which labour he spared her in taking it and +bestowing it among the louers of such varieties, in whose garden it is +yet preserved." Doubtless one of those "lovers" was his friend John +Parkinson, who, in the year 1629, thus wrote concerning it: "One +strawberry more I promised to shew you, which, although it be a wilde +kinde, and of no vse for meate, yet I would not let this discourse passe +without giuing you the knowledge of it. It is in leafe much like vnto +the ordinary, but differeth in that the flower, if it haue any, is +greene, or rather it beareth a small head of greene leaues, many set +thicke together like vnto a double ruffe, in the midst whereof standeth +the fruit, which, when it is ripe, sheweth to be soft and somewhat +reddish, like vnto a strawberry, but with many small harmlesse prickles +on them which may be eaten and chewed in the mouth without any maner of +offence and is somewhat pleasant as a strawberry; it is no great bearer, +but those it doth beare, are set at the toppes of the stalks close +together, pleasant to behold, and fit for a gentlewoman to weare on her +arme, &c., as a rairitie in stead of a flower." + +Merret, in his 'Pinax.' published in 1667, says he found it growing in +the woods of Hyde Park and Hampstead, and Zanoni was the first to figure +it (with the exception of Parkinson's rude woodcut) in his 'Istoria +Botanica,' published in 1675. It is mentioned by Morison and also by +Ray, the latter of whom inserts it in his Synopsis, but without any +habitat; though in his 'Historia Plantarum' he says: "Cantabrigiae in +horto per aliquot annos colui." From this time henceforth the Plymouth +strawberry has become a botanical Dodo, nothing more having been seen or +heard of it except the mere record of the name. In 1766, M. Duchesne +informed the world of the generosity of "M. Monti, Docteur de +Philosophie et de Medecine a Boulogne en Italie," who divided with him a +dried specimen taken from his own herbarium, "Ce present pretieux m'ote +toute incertitude sur la nature de ce Fraisier et sur ses caracteres +monstrueux. Il paroit ne pas avoir aujourd'hui plus d'existence." + +[287] 'Bull. Soc. Bot. France,' 1856, vol. iii, p. 477. + +[288] 'Ann. Sc. Nat.,' 3 ser., vol. ix, p. 86, tabs. v, vi. + +[289] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 695. + +[290] Ibid., vol. iii, 1856, p. 475. + +[291] 'Flora,' 1856, p. 712. + +[292] 'Trans. Linn. Soc.,' vol. xxvi, p. 37. + +[293] 'Bull. Soc. Bot. France,' 1862, vol. ix, p. 36, tab. i, and also +p. 291. + +[294] Ibid., 1857, vol. iv, p. 761. + + + + +CHAPTER II. + +METAMORPHY OF THE FLORAL ORGANS. + + +One of the main arguments adduced by Goethe and others in support of the +now generally received doctrine of the essential morphological identity +of the various whorls of the flower is derived from the frequent +appearance of one organ in the guise of another. The several parts of +the flower become, as it is said, metamorphosed; sometimes the change +is complete, while at other times there may be every conceivable +intermediate condition between one form and another. The sense in which +the terms metamorphosis, substitution, transformation, and the like, are +herein used has already been explained. For the convenience of +arrangement, metamorphosis of the parts of the flower may be divided +into several subdivisions, according to the particular organ affected, +and according to the special kind or degree of change manifested, the +main subdivisions being here classed as Sepalody, Petalody, Staminody, +and Pistillody. + +=Sepalody of the petals.=--This change, spoken of by most authors as +retrograde metamorphosis of the petals into sepals, or as a substitution +of sepals for petals, is obviously a condition that is in most cases +hardly distinguishable from virescence of the corolla, or from +multiplication of the sepals. Nor is this of much consequence unless +there are some special structural features which render the +discrimination a matter of importance, in which case the difficulty is +generally easily surmounted. The flower of the Saint-Valery Apple may +perhaps be cited under this head. In the flower in question there are +neither stamens nor petals, unless the second or inner of sepals be +considered as sepaloid petals (fig. 152). + +[Illustration: FIG. 152.--Flower of St. Valery apple, with sepaloid +petals.] + +M. Alph. de Candolle[295] describes an instance in _Primula Auricula_ in +which the corolla had assumed the appearance of the calyx, but neither +calyx nor corolla in this case possessed perfect stomata. + +This malformation is much less common than the converse one of +calycanthemy. Many of the recorded instances of so-called metamorphosis +of the parts of the flower to sepals have occurred in monocotyledonous +plants, or others in which the calyx and corolla are of the same colour, +and constitute what is frequently termed the perianth; and as this is +usually brightly coloured (not green) it is more convenient to group the +metamorphoses in question under the general term Petalody, which thus +includes all those cases in which the organs of the flower appear in the +form of coloured petal-like organs, whether they be true petals or +segments of a coloured perianth. As the morphological difference between +the organs is one of position merely, there is little objection to be +raised to this course, the less so as the term petalody merely conveys +an idea of resemblance and not of absolute identity. + +Petaloid coloration of the ordinary leaves, or of the bracts, is +mentioned under the chapter relating to colour. + +=Petalody of the calyx--Calycanthemy.=--As with the bracts, so the calyx +in certain instances is naturally coloured, as in _Delphinium_, +_Tropaeolum_, and others. In _Mussaenda_, _Calycophyllum_, _Usteria_, &c., +one or more of the calyx lobes become enlarged normally. Considered +teratologically, petaloid coloration of the sepals is either general or +partial; in the latter case the nerves retain their green colour +longest. There is in cultivation a variety of the primrose called +_Primula calycanthema_, in which the upper part of the calyx becomes +coloured, so that the flower seems to have two corollas placed one +within the other; a similar thing happens in _Mimulus_, in which plant, +as the calyx is permanent while the corolla is deciduous, the coloured +calyx is a great advantage in a horticultural point of view. Morren[296] +says that in order to produce the fine colour of the calyx of _Primula +officinalis_ (var. _smaragdina_) the Belgian gardeners cut away the +corolla in a very early stage, and that in consequence the colouring +matter proper to the corolla is developed in the tube of the calyx, the +edges of the limb remaining green, the middle of the limb being purple +(_Primula tricolor_). + +[Illustration: FIG. 153.--Flower of _Mimulus_, with petaloid calyx.] + +Under this head may be mentioned the occurrence of tubular sepals in +place of the ordinary flat ones in _Helleborus olympicus_; only two of +the sepals were thus affected in a specimen recently observed--a third +exhibited an intermediate condition. + +The normal coloration of the calyx occurs most frequently in +polysepalous calyces; teratological coloration, on the other hand, +occurs especially in gamosepalous flowers. This assertion is borne out +by the frequency of the change in the plants already mentioned, and also +in the following:--_Campanula persicifolia_, _Anagallis arvensis_, +_Gloxinia_, _Syringa persica_,[297] _Calceolaria_, _&c. &c_. In the +last-named plant one or more of the lobes of the calyx may frequently be +seen replaced by a slipper-like petal. + +Among polysepalous plants petaloid sepals have been observed in +_Ranunculus auricomus_, _Rubus caesius_, _&c._ Fleischer also describes a +case of this kind in _Carum carui_.[298] + +It will be seen from the above that in the majority of cases there is no +real metamorphosis or substitution of petal for calyx, but simply an +alteration in colour; nevertheless, a change in form may accompany a +change of colour: this happens especially if there has been any +displacement of organs. Thus, if, in an orchidaceous plant, a sepal be +displaced from any cause, or a petal be twisted out of its natural +position to occupy the place of an absent sepal, that petal will be +sepal-like in form, and _vice versa_. + +=Petalody of the stamens.=--A petaloid condition of the stamens is one +of the commonest of all malformations. A large number of so-called +double flowers (flores pleni)[299] owe their peculiar appearance to this +circumstance. + +It is necessary to distinguish carefully this petaloid development of +the stamens from the corresponding condition of the pistils, and from +that kind of doubling which is a result of multiplication of the +corolla, as in _Datura_, _Campanula_, _Primula_, &c. (flores duplices, +triplices, &c.), or from that produced by true median prolification +(flores geminati, &c.). + +In cases of true petaloid development of the stamens there are usually +numerous intermediate forms between that of the true petals and that of +the perfect stamens; indeed, in _Nymphaea_, _Canna_, and in some other +plants, such a transition occurs normally. Petalody of the stamens may +occur either without material change in the flower or it may exist in +combination or in conjunction with an increased development of parts +(Multiplication), or with a similar change in the carpels, and it is +either partial or complete. + +Among the flowers in which petaloid development of the stamens happens +most frequently may be mentioned those in which the calyx is normally +coloured, as in _Nigella damascena_, _Aguilegia_, and _Delphinium_. + +M. Alph. de Candolle, in the 'Neue Denkschriften,' 1841, described and +figured a singular form of _Viola odorata_, known under the name of +"Bruneau," in Switzerland, in which the stamens are absent, and their +place supplied by a second row of petals, within which is a third series +of petals, representing, says M. de Candolle, the inner row of stamens +that theory suggests should exist in the natural condition. Moreover, +the carpels in this variety are five in number instead of three. In +_Erica Tetralix_ the corolla may not unfrequently be found divided to +the base into its constituent petals, and the place of the stamens +occupied by a series of petal-like structures entirely destitute of +anther. + +In monocotyledonous flowers, especially those with a coloured perianth, +the substitution of segments of the perianth for stamens occurs not +unfrequently. M. Seringe has observed this in the stamens of _Lilium +Martagon_, and there is in cultivation a variety of the white lily, +_Lilium candidum_, sometimes called the double white lily, in which the +segments of the perianth, in place of being arranged in two rows, are +greatly increased in number, and disposed in a spiral manner. In these +flowers, not only are the stamens and pistils thus modified, but also +the upper leaves of the stem. In so-called double tulips there is +likewise a replacement of stamens by coloured segments of the perianth, +but this happens generally in connection with an increase in the number +of organs. Moquin-Tandon remarks having seen in a garden in the environs +of Montpelier a tulip, the stamens of which showed all possible stages +of transition between the form proper to them and that of the perianth. +The pistil in this case was transformed into several small leaves. +Similar appearances have been observed in Iris, Hyacinths, Narcissus, +Colchicum, and Crocus. M. Fournier[300] describes a flower of _Narcissus +Tazetta_ from within the normal perianth of which sprang a second one, +equally provided with a cup and occupying the space usually filled by +the stamens. Flowers of _Narcissus poeticus_ may also be met with in +which the stamens are replaced by six distinct segments exactly +resembling those of the perianth in miniature.[301] + +[Illustration: FIG. 154.--Double columbine, _Aquilegia_--petalody of the +filament.] + +From an examination of these flowers it becomes evident that +petalification is brought about in different flowers in different ways; +sometimes it is the filament which becomes petaloid, sometimes the +anther-lobes, while at other times it is the connective which assumes +the appearance of petals.[302] For instance, in _Solanum tuberosum_, +_S. Dulcamara_, in _Anagallis_, in _Fuchsia_, and some other plants, the +anther-lobes themselves become petaloid, while the filament remains +unchanged. + +In gardens two distinct varieties of Columbine are cultivated, the one +in which the filaments are dilated into the form of flat petals almost +entirely or quite destitute of anthers, while in the other the filament +is present in its usual form, but the anther is developed in the shape +of a tubular hood or spur. + +De Candolle[303] observes that in the _Ranunculaceae_ the species of +_Clematis_ become double by the expansion of the filament, those of +_Ranunculus_ by the dilatation of the anther, and those of _Helleborus_ +by the petal-like development of both filament and anther. In some cases +even on the same plant all three modifications may be seen, as in +Camellias, some of which may be found with petaloid filaments with +anthers on the top, others with the filaments unchanged, but supporting +petaloid anthers, while in others it is the connective alone which is +petal-like. Where the flower naturally contains a large number of +stamens, as in Mallows, Roses, Magnolias, &c., petaloid expansion of the +filament is most common, though it is by no means confined to such +flowers, the change occurring in _Allamanda cathartica_, _Jasminum +grandiflorum_, and many other flowers with few stamens. A similar change +in the anther and connective takes place more frequently in flowers +where the number of stamens is smaller, but there are of course numerous +exceptions to this rule. + +In those cases where there is more than one row of stamens, the +outermost are most liable to this change: thus in _Saxifraga decipiens_, +as shown by Ch. Morren,[304] the outer series of stamens--those opposite +to the sepals--become first affected, and, at a more advanced stage, the +inner row also; and this is the case in most flowers that have their +stamens in two rows. Occasionally it happens that an outer series of +stamens is abortive, or wholly suppressed, while the inner row becomes +petalodic; this was the case in some flowers of _Lilium auratum_ lately +exhibited by Messrs. Veitch. + +Those flowers in which only a portion of the stamens undergo this change +are called semi-double, while in other cases that will be hereafter +mentioned, not only are the stamens thus rendered petaloid, but their +number is also augmented, as in most double roses, pinks, anemones, +poppies, &c. + +In some double flowers, in which the stamens assume more or less +completely the appearance of petals, a singular appearance is afforded +by the presence of four wing-like processes emanating from the central +filaments, two on each side, so that the arrangement may be compared to +two sheets of paper folded in the centre and adherent in that situation, +though perfectly separate elsewhere, except sometimes at the top, where +they form a sort of hood. This change results from an imperfect petalody +of the anther; the two wings on each side of the central vascular cord +represent the front and back walls of an anther lobe, or rather of that +portion of the anther which, under ordinary circumstances, produces +pollen. In the malformed flowers no pollen is formed, at least in the +more complete states of the malformation, but the walls of the anther +lobe become preternaturally enlarged, and petaloid in texture and +appearance. This change occurs in some semi-double rhododendrons and +azaleas, in crocuses, and in a species of violet found at Mentone by Mr. +J. T. Moggridge. + +There are numerous intermediate forms wherein the wing-like processes +may be traced all the way along the filament till they ultimately lose +themselves in the anther-lobes, with which they become continuous. In +some cases, as in _Crocus_ and _Rhododendron_, this is shown even more +clearly by the existence of two perfect pollen-sacs or quarter-anthers, +the remaining portions being petaloid and continuous with the dilated +filament. Not unfrequently these semi-petaloid stamens adhere to the +fronts of the petals, and then it appears, at a first glance, as if +three organs were stuck together, one in front of another, while in +reality there are but two.[305] (See _ante_, p. 35, fig. 12.) + +[Illustration: FIG. 155.--Four-winged filaments of _Rhododendron_.] + +The change in the anther, above alluded to, must not be mistaken for +that far more common one in which only a small portion of the anther +becomes petaloid, forming a sort of lateral wing or appendage to the +polliniferous portion, as happens normally in _Pterandra_, and is common +in some double fuchsias. In this latter instance there is but a single +wing, and the nature of the case is obvious. + +Double flowers of _Orchidaceae_ generally arise from petalification of +the filaments, with or without other coincident changes. What makes +double flowers in this order the more interesting is the development, in +a petaloid condition, of some or all of those stamens which under +ordinary circumstances are wholly suppressed, so that the morphological +structure of the flower, at first a matter of theory, becomes actually +realised. Fig. 156 is a diagram showing the presence of two additional +labella within the ordinary one in a species of _Catasetum_, and +representing two petaloid stamens, thus evidently completing the outer +staminal whorl, of which there is usually but a single representative +(see Peloria, Multiplication, Prolification). In some of these double +orchids it is, however, necessary not to confound a petaloid condition +of the existing column with the development of usually suppressed +stamens in a petaloid form. Thus, in _Lycaste Skinneri_ the column is +frequently provided with two petal-like wings, which might readily be +supposed to be two stamens of the inner whorl adherent to the column; a +little attention, however, to the relative position of these +adventitious wings is generally sufficient to enable the observer to +ascertain the true nature of the appearance.[306] + +[Illustration: FIG. 156.--Diagram of flower of _Catasetum_, with two +labella.] + +Some forms of duplicate or hose in hose corollas are apparently due, not +so much to the formation of a second corolla within the first, as to the +presence of an inner series of petal-like stamens, which, by their +cohesion, form a second pseudo-corolla within the first. The staminal +nature of this pseudo-corolla is inferred from the occasional presence +of anthers on it.[307] In _Datura fastuosa_, as well as in _Gloxinia_, a +pseudo-corolla of this kind sometimes occurs with the addition of a +series of petaloid stamens attached to its outer surface.[308] + +When the petalody specially affects the anther-lobes, as in _Arbutus_, +_Petunia_, _Fuchsia_, _&c._, the venation of the petal-like portion is +very frequently laminar, thus tending to show that the anther is in +such cases really a modification of the blade of the leaf; but as, on +the other hand, we often find petal-like filaments bearing pollen-sacs +on their sides, it is clear that we must not attribute the formation of +pollen to the blade of the leaf only, but we must admit that it may be +formed in the filament as well.[309] + +[Illustration: FIG. 158.--Portion of a double columbine (_Aquilegia_), +showing petalody of the connective.] + +[Illustration: FIG. 159.--Petaloid stamens, _Hibiscus_.] + +Petalody of the connective is of less frequent occurrence than the +corresponding change in the other portions of the stamen. It may be seen +in some forms of double columbine,[310] in which the connective forms a +tubular petal or nectary, and in double petunias and fuchsias. When it +occurs, the true anther-lobes are usually atrophied, and little or no +pollen is formed. + +An occurrence of this nature in _Tacsonia pinnatistipula_, in +conjunction with the partial detachment of the stamens from the +gynophore, led Karsten to establish a genus which he called +_Poggendorffia_.[311] + +From the subjoined list of genera in which petalody of the stamens, in +some form or other, has been observed, it will be seen that it happens +more often in plants with numerous distinct organs (Polypetalae, +Polyandria, Polygynia, &c.) than in other plants with a smaller number +of parts, and which are more or less adherent one to the other. The +tendency to petalification is, moreover, greater among those plants +which have their floral elements arranged in spiral series, than among +those where the verticillate arrangement exists; and in any given +flower, if the stamens are spirally arranged while the carpels are +grouped in whorls, the former will be more liable to petalody than the +latter, and _vice versa_. It has been before remarked, that this +condition is far more common in plants whose petals, &c., have straight +veins, like those in the sheath of a leaf, than in those the venation of +which is reticulate, as in the blade of the leaf. It must also be +remembered that in the same genus, even in the same species, different +kinds of doubling occur. Familiar illustrations of this are afforded in +the case of anemones, columbines, fuchsias, and other plants. + +The existence of "compound stamens" in some flowers, as pointed out by +Payer, and others, and the researches of Dr. Alexander Dickson, confer +additional importance on the subject of petalody, and necessitate the +examination of double flowers with special reference to these compound +stamens, and to the order of their development.[312] The presence of +these compound stamens affords a satisfactory explanation of the +appearance in some double _Malvaceae_, wherein the tufts of adventitious +petals are very liable to be mistaken for buds, produced by axillary +prolification in the axils of the petals, but which are in reality +compound and petaloid stamens. At other times, however, true axillary +prolification exists in these flowers; but then the supplemental florets +have always a calyx, which is wanting in the other instances. + +Petalody of the stamens has been met with most frequently in the +following genera: + + *Ranunculus! + *Anemone! + *Papaver! + *Clematis! + *Hepatica! + *Ficaria! + Thalictrum. + *Caltha! + *Trollius! + *Nigella! + *Aquilegia! + *Delphinium! + *Adonis! + *Paeonia! + *Nelumbium! + *Nymphaea! + *Berberis! + *Papaver! + *Chelidonium! + Sanguinaria. + Podophyllum. + *Mathiola! + *Cheiranthus! + *Iberis! + *Cardamine! + *Hesperis. + *Barbarea! + *Sinapis! + *Brassica! + *Helianthemum! + *Viola! + *Dianthus! + *Saponaria! + *Lychnis! + *Silene! + *Sagina! + *Hibiscus! + *Althaea! + *Malva! + AEsculus! + *Geranium! + *Pelargonium. + *Tropaeolum! + Oxalis! + *Impatiens! + *Camellia! + Thea! + Trifolium! + Medicago! + *Ulex! + Spartianthus. + Clitoria. + Pisum! + Orobus! + Genista! + Spartium! + Cytisus! + Anthyllis. + Coronilla. + Lotus! + *Rosa! + *Kerria! + *Spiraea! + *Fragaria! + *Potentilla! + *Crataegus! + Cydonia. + *Pyrus! + Eriobotrya! + *Amygdalus! + *Prunus! + *Myrtus! + *Punica! + *Philadelphus! + *Deutzia! + *Fuchsia! + Godetia! + Clarkia! + Portulaca! + Ribes! + Saxifraga! + Daucus. + Ixora. + Serissa! + Gardenia! + Lonicera! + Sambucus. + Viburnum. + Scabiosa. + *Campanula! + Platycodon! + Calluna! + Azalea! + Rhododendron! + *Arbutus! + *Erica! + *Anagallis! + *Primula! + *Jasminum! + Syringa! + *Vinca! + *Nerium! + Allamanda! + Tabernaemontana. + *Calystegia! + Convolvulus! + Ipomoea. + *Datura! + *Petunia! + Solanum! + Orobanche. + Gentiana. + Mimulus. + *Antirrhinum! + Gratiola! + *Digitalis! + *Linaria! + Veronica! + Calceolaria! + Achimenes. + Gloxinia! + Clerodendron! + Bignonia. + Cyclamen! + Mirabilis. + Laurus! + Gladiolus! + Crocus! + Iris! + *Galanthus! + Leucojum! + Sternbergia! + Hippeastrum. + *Narcissus! + *Orchis! + Catasetum! + Hydrocharis. + Asphodelus. + *Tulipa! + Scilla. + *Convallaria! + Fritillaria! + *Lilium! + *Hyacinthus! + *Polianthes! + *Hemerocallis! + *Colchicum! + *Sagittaria! + *Tradescantia! + Commelyna! + Tofieldia. + +=Petalody of the pistils.=--Taken by itself, this is much less common +than the corresponding change in the stamens. It generally affects the +style and stigma only, as happens normally in _Petalostylis_, _Iris_, +&c., but this is by no means always necessarily the case. In some of the +cultivated varieties of _Anemone_ and _Ranunculus_ all the parts of the +flower remain in their normal state, except the pistils, which latter +assume a petaloid appearance. + +Many of the double flowers owe their peculiar appearance to the +combination of the following appearances--a petal-like form of the +stamens, increase in the number of these organs and similar changes +affecting the pistils, and is applied to several distinct conditions. If +in any given flower all the stamens and all the pistils become wholly +petaloid, no pollen is formed, and of course no seeds can be produced, +but this very rarely happens, as usually some pollen is produced, and +some ovules capable of being fertilised are developed. + +In double flowers of _Primula sinensis_ it frequently happens that the +capsule is either partially leafy or partly petal-like; in either case +the fruit is open at the extremity, and often destitute of the style and +stigma. It is, however, doubtful if the ovules can be fertilised in +these flowers. + +The following list comprises the names of those genera in which this +change has been most frequently observed, independently of corresponding +alterations in the stamens, but it is more usual for both sets of organs +to be similarly affected. + + *Ranunculus! + *Anemone! + Nigella. + *Papaver! + *Dianthus! + Saponaria! + Viola! + Camellia! + Alcea. + Hibiscus! + Amygdalus! + Lonicera! + Scabiosa. + AEschynanthus! + Primula! + +=Petalody of the ovules.=--The principal changes which occur in the +ovule have already been alluded to at pp. 262-272; it may here be +stated, however, that the ovules are occasionally represented by small +stalked petal-like structures. This happens with especial frequency +among _Cruciferae_.[313] + +=Petalody of the accessory organs.=--A petaloid condition of the disc, +of the scales, or other excrescences from the axis or from the lateral +portions of the flower, is of frequent occurrence, though it is but +rarely that the change is of any great importance in a morphological +point of view. C. Morren has given the name adenopetaly to a case +wherein one of the glands at the base of the petals in _Lopezia_ was +replaced by a petal.[314] A similar change may be seen in the double +Oleander. + +=Staminody of the bracts.=--An instance of this has been already alluded +to in _Abies excelsa_, as observed by Prof. Dickson, and in which some +of the bracts were seen assuming the form and characteristic of the +stamens see _ante_: p. 192. Signor Licopoli met with a similar +substitution of anthers for bracts in _Melianthus major_.[315] + +=Staminody of the sepals and petals.=--In the first named this is of +very rare occurrence. M. Gris has recorded an instance in _Philadelphus +speciosus_[316] which appears to be the only case on record. The +corresponding change in the case of the petals is far more common. De +Candolle cites in illustration of this occurrence flowers of the common +haricot, in which the alae and carina of the corolla were thus +changed.[317] There is in cultivation a form of _Saxifraga granulata_ +wherein the petals are replaced by stamens, so that there are fifteen +stamens. A similar change has been observed in _Capsella +bursa-pastoris_. + +Cramer figures and describes a stamen occupying the place of a petal in +_Daucus Carota_.[318] Turpin[319] describes a similar occurrence in +_Monarda fistulosa_, in which the lower lip terminated in an anther, but +this may have been a case of adhesion. Moquin cites from Chamisso, +_Digitalis purpurea_, and from Jussieu, _Asphodelus ramosus_, as having +presented this change, and Wiegmann[320] has seen anthers developed on +the awns of _Avena chinensis_. In semi-double flowers of _Ophrys +aranifera_ and _Orchis mascula_, the lateral petals are occasionally +partially antheroid, and others occur in which two of the outer series +of stamens, which are ordinarily suppressed, are present, but in a +petaloid state. Reichenbach[321] figures an illustration of this change, +and also Moggridge.[322] + +=Staminody of the pistils.=--The existence of this change has been +denied by several authors, nevertheless, it is of sufficiently common +occurrence. Alexander Braun notices the transformation of pistils into +stamens in Chives (_Allium Scorodoprasum_), and in which three stamens +appeared in the place of as many pistils, and had extrorse anthers, +while the six normal anthers are introrse. In the horse-radish +(_Armoracia rusticana_), two of the carpels are frequently converted +into stamens, while two other organs absent from the normal flower make +their appearance as carpels. Roeper has observed this phenomenon in +_Euphorbia palustris_,[323] and in _Gentiana campestris_.[324] In these +examples one of the carpels was apparently absent, and its place +supplied by an anther. Roeper has also mentioned a balsam with a +supernumerary stamen occupying exactly the position of a carpel.[325] + +Agardh has observed a similar thing in a hyacinth, one half of the fruit +of which contained seeds, and the other half, anthers. B. Clarke +mentions an instance in _Mathiola incana_ in which the carpels were +disunited, and antheriferous at the margin.[326] + +The passage of pistils to stamens in willows has been frequently +remarked, as in _Salix babylonica_, _silesiaca_, _cinerea_, _Caprea_ and +_nigricans_. One of the most curious illustrations of this +transformation in this genus is given by Henry and Macquart (Erst. +Jahrb. des bot. Vereines am m. et n. Rhein., 1837). In the flowers in +question the series of changes were as follows:--first, the ovary opened +by a slit, and then expanded into a cup; next, anther-cells were +developed on the margin of the cup, with stigmas alternating with them, +the ovules at the same time disappearing; lastly, the margin became +divided, and bore three perfect anthers, which in the more perfect +states were raised on three filaments. + +_Campanula persicifolia_, _C. rapunculoides_, and _C. glomerata_ have +been observed to present an anther surmounting the pistil.[327] Double +tulips often present this change, and a like appearance has been +observed in _Galanthus nivalis_, and _Narcissus Tazetta_. + +Moquin mentions the existence of this condition in a female plant of +maize, some of the pistils of which were wholly or partially converted +into anther-like organs. Mohl has recorded an analogous malformation in +_Chamaerops humilis_, and in which the three carpels were normally +formed, and only differed from natural ovaries in this, that along the +two edges of the ventral suture there was a yellow thickening, which a +cross section of the ovary showed to be an anther-lobe filled with +pollen.[328] + +In _Tofieldia calyculata_ a similar substitution of a stamen for a +carpel has been observed by Klotsch,[329] and Weber[330] gives other +instances in _Prunus_ and _Paeonia_. Corresponding alterations may be met +with in cultivated tulips, in the cowslip and other plants. In most of +the above cases the transmutation has been perfect, but in quite an +equal number of cases a portion only of the carpel is thus changed, +generally the style or the stigma; thus Baillon describes the stigmas of +_Ricinus communis_ as having been in one instance antheriferous.[331] +Moggridge figures a flower of _Ophrys insectifera_ in which the +rostellate process was replaced by an anther.[332] + +Mohl remarks that the change of pistils into stamens is more common in +monocarpellary pistils than it is in those which are made up of several +carpels. It seems clear that in this transformation the lobes of the +anther and the development of pollen have no relation to the production +of ovules. + +=Staminody of the accessory organs of the flower.=--The scales that are +met with in some plants, either as excrescences from the petals, or as +imperfect representatives of stamens or other organs, are occasionally +staminoid; thus the scales of _Saponaria officinalis_, of _Silene_, +_Nerium Oleander_, the rays of _Passiflora_, the corona of _Narcissus_, +have all been observed occasionally to bear anthers.[333] In the case of +_Narcissus_ the loose spongy tissue of the corona seems to have the +nearest analogy to the anther-lobes, while the prolonged connective is +more like the ordinary segments of the perianth in texture. The species +in which this change may most frequently be observed are, _N. poeticus_, +_N. incomparabilis_, and _N. montanus_. + +M. Bureau found in some flowers of _Antirrhinum majus_ two petal-like +bodies standing up in front of, or opposite to the two petals of the +upper lip,[334] and similar developments in which each of the two +adventitious segments are surmounted by an anther may be met with +frequently. It does not follow because these organs bear anthers that +they are morphologically true stamens. They are really scales, &c., +taking on themselves accidentally the characters proper to stamens. + +=Pistillody of the perianth.=--The passage of the segments of the +perianth into carpels has been observed frequently in _Tulipa +Gesneriana_, the change in question being generally attended by a +partial virescence. M. Gay is said by Moquin to have observed a flower +of _Crocus nudiflorus_ in which the segments of the perianth were cleft +and fringed at the same time, so that they presented the appearance of +the stigmas. + +[Illustration: FIG. 160.--Flower of tulip, allowing vertical attachment +of a leaf, and also the existence of ovules on the margins of the +segments of the perianth. Some of the parts are removed.] + +=Pistillody of the sepals.=--In some double flowers of the garden pea +communicated by Mr. Laxton, among other peculiarities was a +supernumerary 5-6-leaved calyx, some of the segments of which were of a +carpellary nature, and bore imperfect ovules on their margins, while at +their extremities they were drawn out into styles.[335] + +=Pistillody of the stamens.=--This change whereby the stamens assume more +or less the appearance of pistils is more commonly met with than is the +metamorphosis of the envelopes of the flower into carpels. In some cases +the whole of the stamen appears to be changed, while in others it is the +filament alone that is altered, the anther being deficient, or +rudimentary; while, in a third class of cases, the filament is +unaffected, and the anther undergoes the change in question. In those +instances in which the filament appears to be the portion most +implicated, it becomes dilated so as to resemble a leaf-sheath rather +than a leaf-stalk, as it does usually. + +One of the most curious cases of this kind is that recorded in the +'Botanical Magazine,' (tab. 5160, f. 4) as having occurred in _Begonia +frigida_ already alluded to, and in which, in the centre of a male +flower, were four free ovoid ovaries alternating with as many stamens. +In the normal flowers of this plant, as is well known, the male flowers +have several stamens, while in the female flowers the ovary is strictly +inferior, so that, in the singular flower just described, the perianth +was inferior instead of being superior, as it is usually. It should be +added also that the perianth in these malformed flowers was precisely +like that which occurs ordinarily in the male flowers. + +[Illustration: FIG. 161.--Supernumerary carpels in the orange, arising +from substitution of pistils for stamens.] + +In some varieties of the orange, called by the French "bigarades +cornues," the thalamus of the flower, which is usually short, and +terminated by a glandular ring-like disc, is prolonged into a little +stalk or gynophore, bearing a ring of supernumerary carpels. These +carpels are isolated one from another, and are formed by the +transformation of the filaments of the stamens.[336] + +The additional carpels in the case of the apple of St. Valery, in which +the petals are of a green colour, like the sepals, are by some +attributed to the transformation of the stamens into carpels. These +adventitious carpels frequently contain imperfect ovules and form a +whorl above the normal ones. (See _Pyrus dioica_ of Willdenow.)[337] A +similar change occasionally happens in the stamens of _Magnolia +fuscata_, while in double tulips this phenomenon is very frequent, and +among them may be found all stages of transition between stamens and +pistils, and many of the parts combining the characters of both.[338] +Dunal and Campdera have described flowers of _Rumex crispus_, with seven +pistils, occupying the place of as many stamens. + +[Illustration: FIG. 162.--Substitution of carpels for stamens in +_Papaver_.] + +In _Papaver bracteatum_ a considerable number of the stamens sometimes +become developed into pistils, especially those which are nearest to +the centre of the flower, and in these flowers the filaments are said to +become the ovaries, while the anthers are curled so as to resemble +stigmas. A similar change is not infrequent _Papaver somniferum_. +Goeppert, who found numerous instances of the kind in a field near +Breslau, says the peculiarity was reproduced by seed for two years in +succession.[339] Wigand ('Flora,' 1856, p. 717) has noticed among other +changes the pistil of _Gentiana Amarella_ bearing two sessile anthers. +_Polemonium caeruleum_ is another plant very subject to this change. +Brongniart[340] describes a flower of this species in which the stamens +were represented by a circle of carpels united to each other so as to +form a sheath around the central ovary. By artificial fertilization M. +Brongniart obtained fertile seeds from the central normal ovary as well +as from the surrounding metamorphosed stamens. + +_Cheiranthus Cheiri_ has long been known as one of the plants most +subject to this anomaly. De Candolle even mentions it in his 'Prodromus' +as a distinct variety, under the name of _gynantherus_. Brongniart (loc. +cit.) thus refers to the _Cheiranthus_:--"Sometimes these six carpellary +leaves are perfectly free, and in this case they spread open, presenting +two rows of ovules along their inner edges, or these edges maybe +soldered together, forming a kind of follicle like that of the +columbine; at other times, these staminal pistils are fused into two +lateral bundles of three in each bundle, or into a single cylinder which +encircles the true pistil. In a third set of cases these outer carpels +are only four in number, two lateral and two antero-posterior, all fused +in such a manner as to form around the normal pistil a prism-shaped +sheath, with four sides presenting four parietal placentae, corresponding +to the lines of junction of the staminal carpels." + +In the accompanying figures (fig. 163, _a-d_) the nature of this change +is illustrated. In some of the specimens it is easy to see that the two +shorter stamens undergo the change into carpels later and less perfectly +than the four longer ones, and not infrequently the outer pair are +altogether absent. In most of the flowers of this variety the petals are +smaller and less perfectly developed than usual.[341] + +[Illustration: FIG. 163.--_Cheiranthus Cheiri_, var. _gynantherus_. _a._ +Sepals and petals removed to show carpellodic stamens. _b._ The same +laid open. _c._ Transverse section. _d._ Plan of flower with four +carpel-like stamens, &c.] + +In _Lilium tigrinum_, some specimens of which were gathered by Mr. J. +Salter, in addition to various degrees of synanthy and other changes, +some of the stamens were developed in the form of carpels, adherent by +their edges so as to form an imperfect tube or sheath around the normal +pistil. Fig. 164 shows one of the intermediate organs from these +flowers, in which half the structure seems devoted to the formation of +ovules, while the other half bears a one-celled anther. Lindley[342] has +also described a case of this kind in a species of _Amaryllis_. + +[Illustration: FIG. 164.--Structure half anther, half carpel, _Lilium_.] + +In _Saxifraga crassifolia_ it sometimes happens that mixed with the +stamens, and originating with them, are a number of distinct and +perfectly formed carpels, wholly separated from the normal carpels, in +the centre of the flower. In this particular instance there is usually +no intermediate condition between the stamen and the pistil. +Guillemin[343] also describes a transformation of the stamens into +carpels in _Euphorbia esula_. + +When the anther is involved it may be only partially so, or almost the +whole organ may be transformed. As instances of very partial change may +be cited the passage of the connective into a stigma in _Thalictrum +minus_, or the passage of the points of the anthers into imperfect +styles in some species of bamboo.[344] + +In _Rosa arvensis_ similar transformations have been observed of a +slightly more complex character than those just mentioned, and passing +into more important changes, especially to the formation of pollen +within ovules, formed on the edges of an open carpellodic anther (see p. +186). + +Mr. Berkeley has recorded an analogous case in a gourd in which the +stamens bore numerous ovules (p. 200), and Baillon describes another +gourd in which certain fleshy appendages surrounding the androecium +were provided with ovules.[345] + +Payer, in his 'Organogenie,' p. 38, mentions a stamen of _Dionaea_ +bearing not only an anther, but likewise an ovule. + +_Sempervivum tectorum_ and _S. montanum_, have long been noticed as +being very prone to present this change. Mohl[346] remarks that, in the +transformation of the stamens to the pistil in the common houseleek, the +filament of the stamen generally preserves its form, the anthers alone +undergoing change. At other times, however, the transformation takes +place at the same time, both in the filament and in the anther. When the +stamens are numerous some of them remain in their normal state, while +others, and especially the inner ones, undergo a change. Sometimes all +the stamens are changed simultaneously, while at other times some of +these organs may be found in which the anther is partially filled with +ovules, and partially with pollen. + +In the accompanying figures (fig. 165, _a-h_) a series of intermediate +stages is shown between the ordinary stamen of _Sempervivum tectorum_ +and the ordinary carpel, from which it will be seen that the filament is +little, if at all, affected, and that in those cases where there is a +combination of the attributes of the stamen and of the pistil in the +same organ the pollen is formed in the upper or inner surface of the +leaf-organ, while the ovules arise from the opposite surface from the +free edge, (_b_, _c_, _d_, _e_, _f_, _g_). + +In a drawing made by the Rev. G. E. Smith of a malformed flower of +_Primula acaulis_, and which the writer has had the opportunity of +examining, the stamens are represented as detached from the corolla, and +their anthers replaced by open carpels, with ovules arising, not only +from their edges, but also from their surfaces, while the apex of the +carpellary leaf was drawn out into a long style, terminated by a +flattened spathulate stigma. + +_Delphinium elatum_ is one of the plants in which this change has been +most frequently noticed.[347] + +[Illustration: Fig. 165.--_Sempervivum tecotorum._ _a._ Normal stamen. +_h._ Normal carpel. _b_, _c_, _e_, _f_, _g_. Structure partly staminal, +partly carpellary. _d._ Transverse section through _c_, showing pollen +internally, ovules externally.] + +In willows the change of pistils into staminal organs has been +frequently observed. In _Salix babylonica_ Prof. Schnizlein has +described various transition stages between the carpels and the stamens, +and in one instance, in addition to this change, a perfect cup-shaped +perianth was present, as happens normally in _Populus_[348]. Mr. Lowe +also records the conversion of stamens into ovaries in _Salix +Andersoniana_, and this by every conceivable intermediate +gradation.[349] + +The following list will serve to show what plants are most subject to +this anomaly. It is difficult to draw any accurate inference from this +enumeration, but attention may be called to the frequency of this +occurrence in certain plants, such as the _Sempervivum_, the wallflower, +the poppy, and the heath. Why these plants should specially be subject +to these changes cannot be at present stated. + +By the student of animal physiology such a change as above +described--equivalent to the substitution of an ovary or a uterus for a +testis--would be looked on as next to impossible; the simpler and less +specialised structure of plants renders such a change in them far more +easy of comprehension. + + Thalictrum minus. + Delphinium elatum. + Magnolia fuscata. + Bocconia cordata. + *Papaver bracteatum! + * somniferum! + nudicaule. + Dionaea muscipula! + Barbarea vulgaris. + *Cheiranthus Cheiri! + Cochlearia Armoracia. + Tropaeolum majus. + Citrus Aurantium. + *Sempervivum tectorum! + montanum. + Begonia frigida! + Cucumis, sp. + Cucurbita Pepo. + Pyrus Malus. + Rosa arvensis! + Saxifraga crassifolia! + Myrtus, sp. + Campanula rapunculoides. + Polemonium caeruleum. + Gentiana Amarella. + *Erica Tetralix. + Stachys germanica. + Primula acaulis. + Rumex crispus. + *Salix, sp. plur.! + Euphorbia esula. + Glochidion. + Asphodelus ramosus. + Amaryllis. + Lilium tigrinum! + longiflorum. + *Tulipa Gesneriana! + var. cult. plurim.! + Hemerocallis. + Zea Mays. + Bambusa, sp. + +=Pistillody of the ovule.=--An instance of this extraordinary +transformation in the carnation, as observed by the Rev. Mr. Berkeley, +is given at p. 268. + +FOOTNOTES: + +[295] 'Neue Denkschrift. Schweiz. Gesellsch.,' band v, p. 9. + +[296] 'Bull. Acad. Belg.,' xix, part 2, p. 93. + +[297] Schlechtendal, 'Linnaea,' ix, p. 737. + +[298] Misbilld., 'Cult. Gewachs.,' p. 32. + +[299] Linn., 'Phil. Botan.,' Sec. 120. + +[300] 'Bull. Soc. Bot. France,' 1859, vol. vi, p. 199. + +[301] Seemann's 'Journal of Botany,' vol. iii, p. 105; also Morren, +'Bull. Acad. Belg.,' vol. xx, part 2, p. 264. + +[302] Morren, 'Bull. Belg.,' xviii, p. 503. + +[303] 'Organ. Veg.,' t. i, p. 513. + +[304] 'Bull. Acad. Roy. Belg.,' tome xvii; and Lobelia, p. 65. + +[305] Masters, "On Double Flowers," 'Rep. Internat. Bot. Congress,' +London, 1866. p. 127. + +[306] See also C. Morren, "Sur les vraies fleurs doubles chez les +Orchidees," 'Bull. Acad. Roy. Belg.,' vol. xix, part ii, 1852. p. 171. + +[307] C. Morren, 'Bull. Acad. Belg.,' vol. xx, 1853, part ii, p. 284 +(_Syringa_). + +[308] 'Rep. Bot. Congress,' London, 1866, p. 135, t. vii, f. 14. + +[309] Although it is generally admitted that the filament of the stamen +corresponds to the stalk of the leaf, and the anther to the leaf-blade, +yet there are some points on which uncertainty still rests. One of these +is as to the sutures of the anther. Do these chinks through which the +pollen escapes correspond (as would at first sight seem probable) to the +margins of the antheral leaf, or do they answer to the lines that +separate the two pollen-cavities on each half of the anther one from the +other? Professor Oliver, 'Trans. Linn. Soc.,' vol. xxiii, 1862, p. 423, +in alluding to the views held by others on this subject, concludes, from +an examination of some geranium flowers in which the stamens were more +or less petaloid, that Bischoff's notion as to the sutures of the anther +is correct, viz., that they are the equivalents of the septa of +untransformed tissue between the pollen-sacs. Some double fuchsias +('Gard. Chron.,' 1863, p. 989) add confirmation to this opinion. In +these flowers the petals were present as usual, but the stamens were +more or less petaloid, the filaments were unchanged, but the anthers +existed in the form of a petal-like cup from the centre of which +projected two imperfect pollen-lobes (the other two lobes being +petaloid). Now, in this case, the margins of the anther were coherent to +form the cup, and the pollen was emitted along a line separating the +polliniferous from the petaloid portion of the anther. This view is also +borne out by the double-flowered _Arbutus Unedo_, and also by what +occurs in some double violets, wherein the anther exists in the guise of +a broad lancet-shaped expansion, from the surface of which project four +plates (fig. 157), representing apparently the walls of the pollen-sacs, +but destitute of pollen; the chink left between these plates corresponds +thus to the suture of the normal anther. + +[Illustration: FIG. 157.--Petaloid stamen of _Viola_, with four +projecting plates.] + +The inner or upper portion of the anther-leaf is that which is most +intimately concerned in the formation of pollen; it comparatively rarely +(query ever) happens that the back or lower surface of the antheral leaf +is specially devoted to the formation of pollen. On the other hand, in +cases like those of the common houseleek, where we meet with petaloid +organs combining the attributes of anthers and of carpels, we find the +inner layers devoted to the production of pollen, the outer to the +formation of ovules. + +That the pollen-lobes are not to be taken as halves of a staminal leaf, +but rather as specialised portions of it, not necessarily occupying half +its surface, is shown also in the case of double-flowered _Malvaceae_, in +which the stamens are frequently partly petal-like, partly divided into +numerous separate filaments, each bearing a one-, or it may be even a +two-lobed anther. This circumstance is confirmatory of the opinion held +by Payer, Duchartre, Dickson, and other organogenists, as to the +compound nature of the stamens in these plants. The stamens are here +analogues not of a simple entire leaf, but of a lobed, digitate, or +compound leaf, each subdivision bearing its separate anther. On this +subject the reader may consult M. Mueller's paper on the anther of +_Jatropha Pohliana_, _&c._, referred to at page 255. + +[310] See C. Morren, "On Spur-shaped Nectarines," &c., 'Ann. Nat. +Hist.,' March, 1841, p. 1. tab. 11. + +[311] Karsten, 'Flor. Columb. Spec.,' tab. xxix. + +[312] See Dickson, "On Diplostemonous Flowers," 'Trans. Bot. Soc. +Edin.,' vol. viii, p. 100; and on the Androecium of _Mentzelia_, +_&c_., in Seemann's 'Journal of Botany,' vol. iii, p. 209, and vol. iv +(1866) p. 273 (_Potentilla_, _&c._). + +[313] See Baillon, 'Adansonia,' iii, p. 351, tab. 12, _Sinapis_. + +[314] 'Bull. Acad. Belg.,' xvii, part i, p. 516, c. tab., and +'_Lobelia_,' p. 83. + +[315] Cited in 'Bull. Soc. Bot. France,' xiv, p. 253 ('Rev. Bibl.'). + +[316] 'Bull. Soc. Bot. Fr.,' 1858, p. 331. + +[317] 'Mem. Legum.,' p. 44. + +[318] 'Bildungsabweich, 'Pflanz. Fam.,' tab. 8, f. 12. + +[319] 'Atlas de Goethe' p. 55, t. 4, f. 18. + +[320] Wiegmann, 'Bot. Zeit.,' 1831, p. 5, tab. i. + +[321] 'Ic. Flor. Germ.,' xiii, tab. 112, cccclxiv, f. 2. + +[322] Seemann's 'Journal of Botany,' 1867, p. 317, t. 72, A (_Ophrys_). + +[323] 'Enum. Euphorb.' p. 53. + +[324] 'Linnaea.' i, p. 457. + +[325] 'De Balsam,' p. 17. + +[326] B. Clarke, 'Arrangement of Phaenog. Plants,' p. 23. + +[327] See 'Engelmann,' p. 26, tab. 3, f. 10, 11, 14. + +[328] 'Ann. Sc. Nat.,' ser. 2, t. viii, 1837, p. 58. + +[329] 'Bot. Zeit.,' 4, 1846, 889. + +[330] 'Verhandl. Nat. Hist. Ver. Preuss. Rheinl. und Westph.,' 1858, +1860, p. 381. Cramer also, 'Bildungsabweich,' p. 90, cites a case in +_Paeonia_ where the carpel was open and petaloid, and bore an anther on +one margin, and four ovules on the other. + +[331] 'Euphorbiaceae,' p. 205. + +[332] Seemann's 'Journ. Bot.,' iv, p. 168, tab. 47, f. 1. + +[333] Moquin-Tandon, l. c., 220, _Passiflora_. Masters, 'Journ. Linn. +Soc.,' 1857, p. 159, _Saponaria_. Seemann's 'Journ. Botany,' vol. iii, +p. 107, _Narcissus_. + +[334] 'Bull. Soc. Bot. Fr.,' 1857, p. 452. + +[335] 'Gardeners' Chronicle,' 1866, p. 897. + +[336] Maout, 'Lecons Element.,' vol. ii, p. 488. + +[337] Poiteau and Turpin, 'Arb. Fruit,' t. 37, and Trecul, 'Bull Soc. +Bot. France,' vol. i. p. 307. + +[338] Clos, 'Mem. Acad. Toulouse,' 5 ser., vol. iii. + +[339] 'Bot. Zeit.,' 1850, t. viii, pp. 514, 664. 'Flora,' (B. Z.) 1832, +t. xv, p. 252; also cited in 'Ann. des Serres et des jardins,' vi, pp. +241-5. See also Schlechtendal, 'Bot. Zeit.,' 1845, t. 3, p. 6. + +[340] 'Bull. Soc. Bot. France,' t. viii, p. 453. + +[341] See also Allmann, 'Rep. Brit. Assoc.,' July, 1851. + +[342] 'Theory of Horticulture,' ed. 2, p. 82. + +[343] 'Mem. Soc. Hist. Nat. Paris.' i, 16. + +[344] Gen. Munro, 'Trans. Linn. Soc.,' xxvii, p. 7. + +[345] 'Bull. Soc. Bot. Fr.,' 1857, p. 21. + +[346] 'Ann. Scienc. Nat.,' t. viii, 1837, p. 50, and 'Bot. Zeit.' (R.), +1836, t. xix, p. 513, &c. See also MM. Sourd Dussiples and G. Bergeron, +'Bull. Soc. Bot. France,' viii, p. 349; Von Schmidel, 'Icon. plant. et +Anal. part.' 1782, p. 210, fig. 54. + +[347] Godron, 'Bull. Soc. Bot. Fr.,' xiii, p. 82, Rev. Bibl. + +[348] Cited in Henfrey, 'Bot. Gazette,' iii, p. 12. + +[349] 'Ann. Nat. Hist.,' September, 1856, p. 56. See also Kirschleger, +'Flora (Bot. Zeit.),' xxiv, 1841, p. 340, _Salix alba_. Henschel, 'Flora +(Bot. Zeit.),' 1832, t. xv, p. 253, _S. cinerea_. Hartmann, 'Flora (Bot. +Zeit.),' xxiv, p. 199, _S. nigricans_. Meyer, C. A., 'Bull. Phys. +Math.,' t. x, _S. alba_. + + + + +PART IV. + +HETEROMORPHY. + + +There are certain malformations that have little in common beyond this, +that they cannot readily be allocated in either of the great groups +proposed by writers on teratology. There are also deformities which, +unlike the majority of deviations from the ordinary structure, are +absolute and not relative. While the latter are due to an exaggeration, +or to an imperfection of development, or, it may be, to a partial +perversion in organization, the former differ from the normal standard, +not merely in degree, but absolutely. This is often the case when +disease or injury affects the plant; for instance, in the case of galls +arising from insect-puncture the structure is rather a new growth +altogether, than dependent on mere hypertrophy of the original tissues. +These absolute deformities arising from the causes just mentioned belong +rather to pathology than to teratology strictly so called; but, under +the head of deformities, may be mentioned sundry deviations not +elsewhere alluded to. + + + + +CHAPTER I. + +DEFORMITIES. + + +The special meaning here attached to the term deformity is sufficiently +explained in the preceding paragraph; it remains to give a few +illustrations, and to refer to other headings, such as Heterotaxy, +Hypertrophy, Atrophy, &c., for malformations capable of more rigid +classification than those here alluded to. + +[Illustration: FIG. 166.--Portion of the under surface of a +cabbage-leaf, with horn-like excrescences projecting from it.] + +=Formation of tubes.=--The production of ascidia or pitchers from the +cohesion of the margins of one or more leaves has been already alluded +to (see pp. 21, 30), but there is another class of cases in which the +tubular formation is due, not so much to the union of the margins of a +leaf as to the disproportionate growth of some portions as contrasted +with others, whence arises either a depressed cavity, as in the case of +a leaf, or an expanded and excavated structure, when the stem or some +portion of it is affected. + +The fruit of the rose, the apple, the fig, and many others, is now +generally admitted to be composed externally of the dilated end of the +flower-stalk in which the true carpels become imbedded. Between such +cases and that of a peltate leaf with a depressed centre, such as often +occurs, to some extent, in _Nelumbium_, there is but little difference. + +In cabbages and lettuces there not unfrequently occurs a production of +leaf-like processes projecting from the primary blade at a right angle +(see Enation). Sometimes these are developed in a tubular form, so as to +form a series of little horn-like tubes, or shallow troughs, as in +_Aristolochia sipho_. At other times the nerves or ribs of the leaf +project beyond the blade, and bear at their extremities structures +similar to those just described. + +[Illustration: FIG. 167.--Lettuce leaf, bearing on the back a stalked +cup, arising from the dilatation of the stalk (?).] + +In a variety of _Codiaeum variegatum_ a similar formation may be seen to +a minor extent. Even the common _Scolopendrium vulgare_ occasionally +produces small pitchers of this character, as in the varieties named +_perafero-corautum_, Moore, and _peraferum_, Woll.[350] + +In carnations leaves may sometimes be seen from both surfaces, from +which project long, sharp-pointed tubular spurs at irregular intervals. +A very singular illustration of this is figured by Trattinick,[351], in +which the leaves, epicalyx, sepals, and petals, were all provided with +tubular spurs. + +In _Cephalotus follicularis_ rudimentary or imperfect pitchers may be +frequently met with, in which the stalk of the leaf is tubular and bears +at its extremity a very small rudimentary leaf-blade. It is not in all +cases easy to trace the origin and true nature of the ascidium, as the +venation is sometimes obscure. If there be a single well-marked midrib +the probability is that the case is one of cohesion of the margins of +the leaf; but if the veins are all of about equal size, and radiate from +a common stalk, the pouch-like formation is probably due to dilatation +and hollowing of the petiole. Again, when the result of a union of the +margins of the leaf, the pitcher is generally less regular than when +formed from the hollowed end of a leaf-stalk. Further information is +especially needed as to the mode of development and formation of these +tubular organs, so as to ascertain clearly when they are the result of a +true cupping process, and when of cohesion of the margins of one or more +leaves. (See Cohesion, p. 31. For bibliographical references consult +also A. Braun, 'Flora v. Bot. Zeit.,' 1835, t. xviii, p. 41, +_Aristolochia_.) + +=Tubular formations in the flower.=--A similar formation of tubes +happens in some double flowers; for instance, it is not infrequent in +double flowers of _Primula sinensis_, in which tubular petal-like +structures are attached to the inner surface of the corolla; sometimes +these petaloid tubes replace the stamens, while at other times they +appear to have no relation to those organs. In the particular flowers +now alluded to the tubular form seems due to a dilatation, and not to a +cohesion of the margins. (See Cohesion, p. 23.) These tubular petals +resemble in form and colour almost precisely the normal corolla in +miniature, but are not surrounded by a calyx, nor do they contain +stamens, while the less perfect forms show clearly their origin from a +single tube-like organ. + +[Illustration: FIG. 168.--Corolla of _Primula sinensis_ turned back to +show a tubular petal springing from it. One only is shown for the sake +of clearness; they are generally numerous.] + +The formation of spurs or spur-like tubes in a quasi-regular manner has +been spoken of under the head of Irregular Peloria, p. 228, but we +occasionally meet with tubular processes which seem to occur in an +irregular manner, and to have no reference to the symmetrical plan of +the flower, and which are due probably to the same causes as those which +induce hypertrophy. Such spurs have frequently been seen on the corolla +of _Digitalis purpurea_, _Antirrhinum majus_,[352] _Tulipa Gesneriana_, +and occasionally on the sepals of _Fuchsia_. They are very frequent in +some seasons in the corolla of certain calceolarias (_C. floribunda_). +By Morren this production of adventitious spurs was called +"Ceratomanie." + +[Illustration: FIG. 169.--Corolla of _Calceolaria_, showing irregular +tubular spurs projecting from the lower lip.] + +Similar processes may sometimes be seen in the capsules of _Linaria +vulgaris_, as also in the fruits of some of the Solanums, quite without +reference to the arrangement of the carpels, so that their production +seems to be purely irregular. + +Morren, as previously remarked, gave the name "Solenaidie" to tubular +deformities affecting the stamens, a term which has not been generally +adopted; the deformity in question is by no means of uncommon occurrence +in some double or partially pelorised flowers, as _Antirrhinum_, +_Linaria_, &c. A similar formation of conical out-growths may frequently +be met with in the fruits quite irrespectively of any disjunction of the +carpels. + +=Contortion.=--An irregular twisting or bending of the stem or branches +is by no means of uncommon occurrence, the inducing causes being often +some restriction to growth in certain directions, or the undue or +disproportionate growth in one direction, as contrasted with that in +another. Hence it may arise from insect-puncture, parasitic growth, or +any obstacle to the natural development. Frequently it exists in +conjunction with fasciation, the ends of the branches being curved round +like a shepherd's crook, from the growth on one side being so much +greater than on the other. Sometimes it is a mere exaggeration of a +normal condition; thus, in what are termed flexuose stems the stem +twists alternately to one side or another, frequently in association +with an oblique form of the leaf. This state is sometimes present to an +extreme degree, as in some varieties of shrubs (_Crataegus_, _Robinia_, +&c.) cultivated for their singularly tortuous branches. + +[Illustration: FIG. 170.--Portion of the culm of a _Juncus_, bent +irregularly.] + +[Illustration: FIG. 171.--Portion of a branch of _Crataegus oxyacantha_, +var. _tortuosa_.] + +Such cases as those just mentioned, however, are but slightly irregular +compared to others in which the deformity exists to such an extent that +the traces of the ordinary mode of growth are almost obliterated. M. +Moquin-Tandon[353] alludes to a case of this kind in a species of pine +(_Pinus_), in which a branch ended in four unequal divisions, which were +strongly curved from without inwards, then became united in pairs, these +latter in their turn blending into a single mass. + +In the case of some beeches growing in the forest of Verzy, near Rheims, +the trunks of the trees are contorted in every direction, and, at a +height of from fifteen to twenty feet, a number of branches are also +given off, also much contorted, and occasionally intergrafted, so that +it seems as if a heavy weight had been placed on the trees and literally +flattened them. Similar malformations may occasionally be met with in +the branches of the oak, and commonly in the weeping ash. + +M. Fournier[354] mentions the stems of _Ruscus aculeatus_ rolled in a +circle, others twisted spirally. + +The phenomenon is not confined to woody plants, but has been met with in +chicory, in _Antirrhinum_, and other herbaceous species. + +It is very difficult in some cases to separate these instances of +irregular torsion from those in which the twisting takes place in a more +or less regular spiral direction. In the former case the fibres of the +plant are only indirectly involved, but in the latter the fibres +themselves are coiled spirally from right to left, or _vice versa_ +(spiral torsion), while not unfrequently both conditions may be met with +at the same time. + +The leaves also are subject to similar deformities, of which a notable +illustration has been recorded in the case of the date palm, _Phoenix +dactylifera_, originally observed by Goethe, and figured and described +by Jaeger;[355] the leaves are folded and twisted in every direction, in +consequence of the fibrous band or cord which surrounds the leaves, and +which generally breaks as the leaflets increase in size, remaining from +some cause or other unbroken, and thus serving to restrain the growth. + +A similar irregularity of growth occurs, not unfrequently, in the case +of crocus leaves, when in the course of their growth, as they push their +way through the soil, their progress becomes checked either by a stone +or even by frost. + +=Spiral torsion.=--Growth in a spiral direction, and the arrangement of +the various organs of the plant in a spiral manner, are among the most +common of natural phenomena in plants.[356] Fibres are coiled spirally +in the minute vessels of flowering plants, and are not wholly wanting +even among fungi. The leaf-organs are very generally spirally arranged; +the leaf-stalks are often so twisted as to bring leaves on one plane +which otherwise would occupy several. In the leaf itself we have a +spiral twist taking place constantly in _Alstroemeria_, in _Avena_, and +other plants. A similar tendency is manifested in the flower-stalks, as +in _Cyclamen_ and _Vallisneria_, and the whole inflorescence, as in +_Spiranthes_. Even the bark and wood of trees is often disposed +spirally. This is very noticeable in some firs, and in the bark of the +sweet chestnut (_Castanea_), of _Thuja occidentalis_, and other trees. +The knaurs or excrescences which are sometimes found on the roots or +stems of trees afford other illustrations of this universal tendency. +These bodies consist of a number of embryo buds, which, from some cause +or other, are incapable of lengthening. On examination every rudimentary +or undeveloped bud may be seen to be surrounded by densely crowded +fibres arranged spirally. + +The axes of nearly all twining plants are themselves twisted, and +twisted in a direction corresponding to the spontaneous revolving +movement exhibited by these plants, as in the hop, the convolvulus, +passion flower, &c., the degree of twisting being dependent to a great +extent on the roughness of the surface around which the stem +twines[357]. + +Considered as an exceptional occurrence, it occurs frequently in certain +plants, and, when it affects the stem or branches, necessarily causes +some changes in the arrangement of the parts attached to them; thus, +spiral torsion of the axial organs is generally accompanied by +displacement of the leaves, whorled leaves becoming alternate, and +opposite or whorled leaves becoming arranged on one side of the stem +only. Frequently also this condition is associated with fasciation, or, +at least, with a distended or dilated state. An illustration of this in +_Asparagus_ has been figured at p. 14. + +Very often the leaves are produced in a spiral line round the stem, as +in a specimen of _Dracocephalum speciosum_ described and figured by C. +Morren. The leaves of this plant are naturally rectiserial and +decussate, but, in the twisted stem the leaves were curviserial, and +arranged according to the 5/13 plan. Now, referring to the ordinary +notation of alternate leaves, we shall have the first leaf covered by +the fifth, with two turns of the spiral; since decussate leaves result +from two conjugate lines, the formula will be necessarily 2/5. The +fraction 5/13 hence comes regularly into the 2/5 series (2/5, 3/8, +5/13). Thus, the leaves in assuming a new phyllotaxy, take one quite +analogous to the normal one. + +One of the most curious instances that have fallen under the writer's +own observation occurred in the stem of _Dipsacus fullonum_. (See +'Proceedings of the Linnean Society,' March 6, 1855, vol. ii, p. 370). +The stem was distended, and hollow, and twisted on itself; its fibres, +moreover, were arranged in an oblique or spiral direction; the branches +or leaf-stalks, which usually are arranged in an opposite and decussate +manner, were, in this case, disposed in a linear series, one over the +other, following the line of curvature of the stem. When the course of +the fibres was traced from the base of one of the stalks, upward around +the stem, a spiral was found to be completed at the base of the second +stalk, above that which was made the starting point. Now, if opposite +leaves depend on the shortened condition of the internode between the +two leaves, then, in the teazel-stem just described, each turn of the +spiral would represent a lengthened internode; and, if the fibres of +this specimen could be untwisted, and made to assume the vertical +direction, and, at the same time, the internodes were shortened, the +result would be the opposition of the branches and the decussation of +the pairs; this explanation is borne out by the similar twisting which +takes place so frequently in the species of _Galium_ and other +_Rubiaceae_. + +[Illustration: FIG. 172.--Twisted stem of _Dipsacus fullonum_.] + +G. Franc[358] was one of the first to notice this twisting in _Galium_, +and M. Duchartre,[359] in mentioning a similar instance, gives the +following explanation of the appearance which will be found to apply to +most of these cases. In the normal stem of _Galium Mollugo_ the +branches are opposite in each verticil and crossed in the two +successive ones. The stem is four-angled, each angle having a nerve. +Each of these nerves, springing from the origin of a branch in one +whorl, terminates in the interval which separates the point of origin of +the two branches in the whorl next above it. In the deformed stem one of +the nerves corresponds to the insertion of a branch, its neighbour is in +the adjoining vacant space; hence it results that four nerves correspond +to two branches and to two consecutive interspaces, and hence the +analogy between a single normal internode provided with its two branches +and its four nerves. What confirms this inference is that the nerve, +which begins at the point of origin of a branch, after making one spiral +turn round the stem, terminates in the interval that separates the two +following branches, just as in a branch of the normal stem it ends in +the upper whorl between the two next branches. The torsion, then, in +this _Galium_ caused the separation of the two opposite branches of the +same verticil, and placed them one above another, and this being +reproduced in all the whorls, all the branches come to be arranged on +the same longitudinal line. The leaves are susceptible of the same +explanation; they are inserted in groups of three or four in one arc +round the origin of each branch. In the malformation each series or +group of four leaves, with its central branch, is equivalent to half a +whorl of the natural plant with its axillary branch. In other words, the +malformation consists in a torsion of the stem, which separates each +whorl into two distinct halves; these half-whorls, with their axillary +branches, are placed on a single longitudinal series one above another. +This case is quoted at some length, as it is an admirable example of a +very common form of malformation in these plants. + +In some parts of Holland where madder is cultivated a similar +deformation is particularly frequent. The leaves, however, are not +always grouped in the way in which they were described by M. Duchartre, +but more commonly form a single continuous line; when arranged in +leaf-whorls it generally happens that some of the leaves are turned +downwards, while others are erect. It has been said that this condition +occurs particularly frequently in plants growing in damp places. It is +certainly true that spiral torsion of the stem is specially frequent in +the species of _Equisetum_, most of which grow in such spots. In these +plants either the whole of the upper part of the stem is thus twisted, +or a portion only: thus Reinsch[360] cites a case in _Equisetum +Telmateia_, where the upper and lower portions of the stem were normal, +while the intermediate portion was twisted spirally. In this instance +the whorl next beneath the spiral had twenty-eight branchlets, and that +immediately above it thirty. Along the course of the spire there were +two hundred and three; dividing this latter number by the mean of the +two preceding, it was seen that the spire included the constituents of +seven ordinary verticils. + +[Illustration: FIG. 173.--Stem of _Galium_ spirally twisted. From a +specimen communicated by Mr. Darwin.] + +Here also may be mentioned a curious bamboo, the stem of which is +preserved in the British Museum, and in which the internodes, on the +exterior, and the corresponding diaphragms and cavities within are +spiral or oblique in direction. + +The root is also subject to the same malformation, the inducing cause +being usually some obstruction to downward growth, as when a plant has +been grown in a small pot, and becomes, as gardeners say, pot-bound. + +[Illustration: FIG. 174.--Showing "pot-bound" root twisted spirally +(from the 'Gard. Chron.,' 1849).] + +The axial portion of the flower, the thalamus, is also occasionally +twisted in a spiral direction, the lateral parts of the flower being in +consequence displaced. Morren spoke of this displacement of the floral +organs as "speiranthie."[361] + +Morren draws a distinction between spiral-torsion or spiralism and the +less regular torsion spoken of in the preceding section; in the former +case not only is the axis twisted, but its constituent fibres also. The +condition in question in some cases seems to be inherited in the +seedling plants. + +The following is a list of the plants in which spiral torsion of the +stem or branches has been most frequently observed. (See also under +Fasciation and Contortion.) + + Hesperis matronalis. + Dianthus barbatus. + Pyrus Malus. + torminalis. + Cercis siliquastrum! + Punica Granatum. + Robinia pseudacacia! + Rubia tinctorum. + Dipsacus fullonum! + pilosus. + Gmelini. + Scabiosa arvensis. + *Valeriana officinalis! + dioica! + Galium aparine! + * Mollugo! + verum! + Hippuris vulgaris! + Veronica spicata. + longifolia. + Hyssopus officinalis. + Thymus Serpyllum. + Lamium purpureum! + Dracocephalum speciosum. + Mentha aquatica. + Mentha viridis. + Fraxinus vulgaris! + Sambucus nigra. + Zinnia. + Phylica. + Beta. + Rumex, sp. + Ulmus campestris. + Casuarina rigida. + Abies excelsa! + Lilium Martagon! + candidum. + *Asparagus officinalis! + Sagittaria sagittifolia. + Epipactis palustris. + Triticum repens! + Lolium perenne! + Phleum pratense. + Juncus conglomeratus! + Scirpus lacustris. + Equisetum Telmateia. + limosum. + fluviatile. + arvense! + +Among the more important papers relating to this subject may be +mentioned: + + Moquin-Tandon, 'El. Ter. Veg.,' p. 181. Kros, 'De Spira in + plantis conspicua.' Morren, 'Bull. Acad. Roy. Belg.,' 1851, + tom. xviii, part i, p. 27. Milde, 'Nov. Act. Acad. Leop. Carol. + Nat. Cur., 1839. Ibid., vol. xxvi, part ii, p. 429, + _Equisetum_. Irmisch, 'Flora,' 1858, t. ii, _Equisetum_. + Vrolik, 'Nouv. Mem. Instit. Amsterdam,' _Lilium_. + Schlechtendal, 'Bot. Zeit.,' xiv, p. 69, et v, p. 66. De + Candolle, 'Organ. Veget., t. i, p. 155, tab. xxxvi, _Mentha_, + _&c._ Alph. de Candolle, 'Neue Denkschr. Allg. Schweiz. + Gesellschft.,' band v, tab. vi, _Valeriana_. Duchartre, 'Ann. + Sc. Nat.,' ser. 3, vol. i, p. 292. 'Gardeners' Chronicle,' July + 5, 1856, p. 452, _c. ic. xylogr._, spiral branches from + Guatemala--tree not known. + +=Spiral twisting of the leaf= is scarcely of so common occurrence as the +corresponding condition in the stem. In _Alstroemeria_ it occurs +normally, as also in some grasses. In the variety _annularis_ of _Salix +babylonica_ the leaf is constantly coiled round spirally. A similar +contortion occurs in a variety of _Codiaeum variegatum_ lately introduced +from the islands of the South Seas by Mr. J. G. Veitch. + +Fern fronds are occasionally found twisted in the same manner, _e.g._ +_Scolopendrium vulgare_ var. _spirale_.[362] + +=Adventitious tendrils.=--Under ordinary circumstances tendrils may be +described as modifications of the leaf, the stipule, the branch, or of +the flower stalk, so that it is not a matter of surprise to find +tendrils occasionally springing from the sepals or petals, as indeed +happens normally in _Hodgsonia_, _Strophanthus_, _&c._ + +M. Decaisne[363] found a flower of the melon in which one of the +segments of the calyx was prolonged into a tendril, and Kirschleger +records a similar instance in the cucumber, while Mr. Holland ('Science +Gossip,' 1865, p. 105) mentions a case in which one of the prickles on +the fruit of a cucumber had grown out into a tendril. + +In _Cobaea scandens_ the foliar nature of the tendril is shown by the +occasional presence of a small leaflet on one of the branches of the +tendril, and a similar appearance may frequently be seen in +_Eccremocarpus scaber_. On the other hand, in the vine, the axial nature +of the tendril is revealed by the not infrequent presence of flowers or +berries on them, as also in _Modecca_ and some _Passifloraceae_. + +Darwin, speaking of the tendrils of _Bignonia capreolata_, says it is a +highly remarkable fact that a leaf should be metamorphosed into a +branched organ, which turns from the light, and which can, by its +extremities, either crawl like a root into crevices, or seize hold of +minute projecting points, these extremities subsequently forming +cellular masses, which envelope by their growth the first fibres and +secrete an adhesive cement. + +=Interrupted growth.=--This term is here used in the same sense as in +ordinary descriptive botany, as when an "interruptedly pinnate" leaf is +spoken of. A similar alternation may be observed occasionally as a +teratological occurrence, though it is not easy to account for it. + +[Illustration: FIG. 175.--Interrupted growth of Radish (from the +'American Agriculturist.')] + +[Illustration: FIG. 176.--Interrupted growth in Apple.] + +Fig. 175 shows an instance of the kind in a radish, and fig. 176 a +similar deformity in the case of an apple, the dilatation of the +flower-stalk below the ordinary fruit producing an appearance as if +there were two fruits one above another. + +In leaves this peculiar irregularity of development is more common. + +In some varieties of _Codiaeum variegatum_ the leaves resemble those of +_Nepenthes_, as the basal portion is broad, and terminates in a +projecting midrib destitute of cellular covering, and this again +terminates in a small pouch or pitcher. Somewhat similar variations may +be found in ferns, especially _Scolopendrium vulgare_. + +Instead of the pouch there is formed sometimes in the plant last +mentioned a supplementary four-lobed lamina, the four lobes being in two +different planes, and diverging from the midrib, so that the section +would resemble [Symbol: Sideways X], the point of intersection of the x +representing the position of the midrib. This four-winged lamina is thus +very similar to the four-winged filaments described and figured at p. +289, and to the leaf-like anther of _Jatropha_ described by M. Mueller, +p. 255. + +=Cornute leaves= (_Folia cornuta_).--The condition to which this term +applies is that in which the midrib, after running for a certain +distance, generally nearly to the point of the leaf, suddenly projects, +often in a plane different from that of the leaf, and thus forms a small +spine-like out-growth. Should this happen to be terminated by a second +laminar portion, an interrupted leaf would be formed. In _Scolopendrium +vulgare_ and other ferns this condition has been noticed, as also in +some of the varieties of _Codiaeum variegatum_ already referred to. + +=Flattening.=--There are some plants whose stem or branches, instead of +assuming the ordinary cylindrical form, are compressed or flattened; +such are some species of _Epiphyllum_, _Coccoloba_, _Bauhinia_, &c. The +same thing occurs in the leaf-like branches of _Ruscus_, the +flower-stalks of _Xylophylla_, _Phyllanthus_, _Pterisanthes_. Martins +proposes to apply the word 'cladodium' to such expansions, just as the +term phyllodium is applied to the similar dilatation of the leaf-stalks. +If we exclude instances of fasciation, _i.e._ where several branches +are fused together and flattened, we must admit that this flattening +does not occur very often as a teratological appearance. + +Mr. Rennie figures and describes a root of a tree which had become +greatly flattened in its passage between the stones at the bottom of a +stream, and had become, as it were, moulded to the stones with which it +came into contact.[364] + +The spadix of _Arum_, as also of the cocoa-nut palm, has been observed +flattened out, apparently without increase in the number of organs. + +When the blade of the leaf is suppressed it often happens that the stalk +of the leaf is flattened, as it were, by compensation, and the petiole +has then much the appearance of a flat ribbon (phyllode). This happens +constantly in certain species of _Acacia_, _Oxalis_, &c., and has been +attributed, but doubtless erroneously, to the fusion of the leaflets in +an early state of development and in the position of rest.[365] + +In some water plants, as _Sagittaria_, _Alisma_, _Potamogeton_, &c., the +leaf-stalks are apt to get flattened out into ribbon-like bodies; and +Olivier has figured and described a _Cyclamen_, called by him _C. +linearifolium_, in which, owing to the suppression of the lamina, the +petiole had become dilated into a ribbon-like expansion--deformation +rubanee of Moquin. + +FOOTNOTES: + +[350] Moore, 'Nature Printed Ferns,' 8vo edition, vol. ii, p. 154, et p. +173. + +[351] 'Flora (B. Z.),' 1821, vol. iv, p. 717, c. tab. + +[352] Chavannes, 'Mon. Antirrh.' + +[353] 'Bull. Soc. Bot. France,' t. vii, 1860, p. 877. + +[354] Ibid., t. iv, 1857, p. 759. + +[355] Jaeger, "De monstrosa folii _Phoenicis dactyliferae_ +conformatione a Goetheo olim observata," 'Act. Acad. Leop. Car. Nat. +Cur.,' vol. xvii, suppl., p. 293, c. tab. color. iv. + +[356] See Goethe, 'Ueber die spiral Tendenz.' + +[357] See Darwin "On Climbing Plants," 'Journ. Linn. Soc. Botany,' vol. +ix, p. 5. + +[358] 'Ephem. Nat. Cur.,' dec. 2, ann. 1, 1683, p. 68, fig. 14. + +[359] 'Ann. des Scienc. Nat.,' third series, vol. i, 1844, p. 292. + +[360] 'Flora' Feb. 4, 1858, p. 69, tab. ii, f. 3, and also 'Flora,' +1860, p. 737, tab. vii, f. 9. + +[361] 'Bull. Acad, Belg.,' t. xvii, p. 196, "Lobelia," p. 53, c. tab. + +[362] Moore, 'Nature-printed Ferns,' 8vo edition, vol. ii, p. 183. + +[363] 'Bull. Soc. Bot. Fr.,' 1860, vol. vii, p. 461. See also Naudin, +'Ann. Sc. Nat.,' 4 ser., t. iv, p. 5. Clos, 'Bull. Soc. Bot. Fr.,' t. +iii, p. 546. + +[364] London's 'Magazine Nat. Hist.,' vol. ii, p. 463. + +[365] C. Morren, 'Bull. Acad. Belg.,' 1852, t. xix, part iii, p. 444. + + + + +CHAPTER II. + +POLYMORPHY. + + +Usually the several organs of the same individual plant do not differ to +any great extent one from another. One adult leaf has nearly the same +appearance and dimensions as another; one flower resembles very closely +another flower of the same age and so on. Nevertheless it occasionally +happens that there is a very considerable difference in form in the same +organs, not only at different times, but it may also be at the same +time. Descriptive botanists recognise this occurrence in the case of +leaves, and apply the epithet heterophyllous to plants possessed of +these variable foliar characters. In the case of the flower, where +similar diversity of form occasionally exists, the term dimorphism is +used. + +As these phenomena appear constantly in particular plants, they are +hardly to be looked on, under such circumstances, as abnormal, but where +they occur in plants not usually polymorphic, they may be considered as +coming within the scope of teratology. + +=Heterophylly.=--As a general rule, the leaves or leaf-organs in each +portion of a plant, from the rhizome or underground axis, where it +exists, to the carpellary leaf, have their own special configuration, +subject only to slight variations, dependent upon age, conditions of +growth, &c. The cotyledons are very uniform in shape in each plant, and +are scarcely ever subject to variation. The leaves near the base of the +stem, the root-leaves as they are not unfrequently called, sometimes +differ in form from the stem-leaves; these again differ from the bracts +or leaves in proximity to the flower. The floral envelopes themselves, +as well as the bud-scales, all have their own allotted form in +particular plants, a form by which they may, in most cases, be readily +recognised. Hence, then, in the majority of plants there is naturally +very considerable difference in the form of the leaf-organs, according +to the place they occupy and the functions they have to fulfil; but, in +addition to this, it not unfrequently happens that the leaf-organs in +the same portion of the stem are subject to great variation in form. +This is the condition to which the term heterophylly properly applies. +The variation in form is usually dependent on a greater or less degree +of lobing of the margin of the leaf; thus, in the yellow jasmine, almost +every intermediate stage may be traced from an ovate entire leaf to one +very deeply and irregularly stalked. _Broussonettia papyrifera_, and +_Laurus Sassafras_, and the species of _Panax_, may be mentioned as +presenting this condition. Sometimes in the last-named genus, as also in +_Pteridophyllum_, every gradation between simple and compound leaves may +be traced. The horse-radish (_Cochlearia Armoracia_) may also be +instanced as a common illustration of polymorphism in the leaves. In +ferns it is likewise of frequent occurrence, markedly so in +_Scolopendrium D'Urvillei_, in which plant every gradation from a simple +oblong frond to an exceedingly divided one may be found springing from +the same rhizome at the same time. + +[Illustration: FIG. 177.--_Syringa persica laciniata_, showing +polymorphous leaves.] + +A similar protean state, but little less remarkable, occurs in many of +our British ferns, notably in _Scolopendrium vulgare_, of which Mr. +Moore enumerates no fewer than 155 varieties,[366] many of the forms +occurring on the same plant at the same time. Cultivators have availed +themselves of this tendency to produce multiform foliage, not only for +the purposes of decoration or curiosity, as in the many cut-leaved or +crisped-leaved varieties, but also for more material uses, as, for +instance, the many varieties of cabbages, of lettuces, &c. Most of these +variations are mentioned under the head of the particular morphological +change of which they are illustrations. + +The effect of a change in the conditions of growth in producing +diversity in the form of the leaf may be here alluded to. _Ficus +stipulata_, a plant used to cover the walls of plant-stoves in this +country, and growing naturally on walls in India, like ivy, produces +leaves of very different form, size, and texture, when grown as a +standard, from what it does when adhering to a wall. _Marcgraavia +umbellata_ furnishes another example of a similar nature, as indeed, to +a less extent, does the common ivy. + +Allusion has been already made to the occasional persistence of forms in +adult life, which are commonly confined to a young state, as in the case +of some conifers which present on the same plant, at the same time, two +different forms of leaves. Mention has also been made of the presence of +adventitious buds on leaves and in other situations. The leaves that +spring from these buds are usually of the same form as the other leaves +of the plant, but now and then they differ. Of this a remarkable +illustration is afforded by a fern, _Pteris quadriaurita_, in which the +fronds emerging from an adventitious bud are very different from the +ordinary fronds. + +[Illustration: FIG. 178.--Portion of a frond of _Pteris quadriaurita_, +with an adventitious bud, the form of the constituent foliage of which +is very different from that of the parent frond.] + +=Dimorphism.=--This term, applied specially to the varied form which the +flowers or some of their constituent elements assume on the same plant, +is an analogous phenomenon to what has been above spoken of as +heterophylly, and, like it, it cannot, except under special +circumstances, be considered as of teratological importance. A few +illustrative cases, however, may here be cited. + +Sir George Mackenzie describes a variety of the potato[367] (_Solanum +tuberosum_), which produces first double and sterile flowers, and +subsequently single fertile ones; the other portions of the plant do not +differ much. + +_Stackhousia juncea_, according to Clarke, has mixed with its perfect +flowers a number of apetalous blossoms destitute of anthers.[368] + +This peculiarity is well exemplified in the tribe _Gaudichaudieae_ of the +order _Malpighiaceae_. A. de Jussieu, in his monograph, speaks of these +flowers as being very small, green, destitute of petals, or nearly so, +with a single, generally imperfect anther; the carpels also are more or +less imperfect, but not sufficiently so to prevent some seeds from being +formed. A similar production of imperfect flowers has been noticed in +many other orders, _e.g._ _Violaceae_, _Campanulaceae_, &c. In some cases +these supplementary blossoms are more fertile and prolific in good seeds +than are the normally constructed flowers. M. Durieu de Maisonneuve +alludes to a case where flowers of this description are produced below +the surface of the ground. The plant in question is _Scrophularia +arguta_, and it appears that towards the end of the summer the lowest +branches springing from the stem bend downwards, and penetrate the soil; +the branches immediately above the lowest ones also bend downwards, but +do not always enter the earth. These branches bear fertile flowers: +those which are completely below the soil are completely destitute of +petals; those which are on the surface have a four-lobed corolla whose +divisions are nearly equal, like those of _Veronica_.[369] + +To Sprengel, and specially to Darwin, physiologists are indebted for the +demonstration of the relation of di- and trimorphic flowers to +fertilisation. In certain genera of orchids, such as _Catasetum_, &c., +flowers of such different form are produced that botanists, without +hesitation, considered them as belonging to different genera, until the +fact of their occasional production on the same plant showed that they +were not of even specific importance. It was reserved for Mr. Darwin to +show experimentally that these very different flowers are really sexual +forms of one and the same species, ordinarily occurring on different +plants, i.e. dioecious, but occasionally formed on the same spike. The +same excellent observer has demonstrated that the di- and trimorphic +forms of _Primula_, of _Linum_, _Lythrum_, and other plants--forms +differing mainly in the relative length of the stamens and styles, are +also connected with striking differences in the number of perfect seeds +produced. The most perfect degree of fertility is obtained when the +stigma of one form is fertilised by the pollen taken from stamens of a +corresponding height. On the other hand, when the union is, as Mr. +Darwin states, illegitimate, that is, when the pollen is taken from +stamens not corresponding in length to the style, more or less complete +sterility ensues in the progeny, sometimes even utter infertility, such +as happens when two distinct species are crossed, so that, in point of +fact, the offspring of these illegitimate unions correspond almost +precisely to hybrids.[370] + +Mere variations of form arising from hybridisation or other causes +hardly fall within the limits of this work, though it is quite +impossible to say where variations end and malformations begin. There +are, however, two or three cases cited by Mr. Darwin[371] from Gallesio +and Risso to which it is desirable to allude. Gallesio impregnated an +orange with pollen from a lemon, and the fruit borne on the mother tree +had a raised stripe of peel like that of a lemon both in colour and +taste, but the pulp was like that of an orange, and included only +imperfect seeds. Risso describes a variety of the common orange which +produces "rounded-oval leaves, spotted with yellow, borne on petioles, +with heart-shaped wings; when these leaves fall off they are succeeded +by longer and narrower leaves, with undulated margins, of a pale green +colour, embroidered with yellow, borne on foot-stalks without wings. +The fruit whilst young is pear-shaped, yellow, longitudinally striated +and sweet; but, as it ripens, it becomes spherical, of a reddish-yellow, +and bitter." + +=Sports or bud variations.=--These curious departures from the normal +form can only be mentioned incidentally in this place, as they pertain +more to variation than to malformation. + +The occasional production of shoots bearing leaves, flowers, or fruits +of a different character from those found on the normal plant, is a fact +of which gardeners have largely availed themselves in the cultivation of +new varieties. The productions in question have been attributed to +various causes, such as cross-breeding, grafting, budding, dissociation +of hybrid characters, or reversion to some ancestral form, all of which +explanations may be true in certain cases, but none of them supply the +clue to the reason why one particular branch should be so affected, and +the rest not; or why the same plant, at the same time, as often happens +in Pelargoniums, should produce two, three, or more "sports" of a +different character. + +These bud variations may be perpetuated by grafts or by cuttings, +sometimes even by seed. With reference to cuttings a curious +circumstance has been observed, viz., that if taken from the lower part +of the stem, near the root, the peculiarity is not transmitted, but the +young plant reverts to the characters of the typical form (Carriere). +This circumstance, however, is not of universal occurrence. + +For further particulars on this interesting subject the reader is +referred to Darwin's 'Variation of Animals and Plants,' i, p. 373, where +numerous references are given, and wherein certain well-known and highly +remarkable instances, such as the _Cytisus Adami_, the trifacial orange, +&c., are discussed. + +FOOTNOTES: + +[366] 'Nature-printed Ferns,' 8vo edition, vol. ii, p. 197. + +[367] 'Gard. Chron.,' 1845. p. 790. + +[368] 'A New Arrangement of Phaenog. Plants,' p. 36. + +[369] 'Bull. Soc. Bot. France,' 1856, t. iii, p. 569. + +[370] The reader will find an abstract of Mr. Darwin's views in his work +on the 'Variation of Animals and Plants,' vol. ii, p. 181. + +[371] Loc. cit., i, 336. + + + + +CHAPTER III. + +ALTERATIONS OF COLOUR.[372] + + +Changes in the colour of the several organs of plants are more often +either pathological or the result of variation than of malformation +properly so called. + +Alterations in colour arise from a diminished or an increased amount of +colouring matter, or from an unusual distribution of the solid or fluid +matters on which the colour depends. The superposition of cells +containing colouring material of different tints produces naturally a +very different set of hues from those which are manifested when the +colours are not blended. Referring the reader to the ordinary text-books +on vegetable physiology and chemistry for details as to the nature and +disposition of colouring materials in plants under natural +circumstances, it will only be necessary to cite a few instances of +deviation from the general colour of plants or their organs. + +=Albinism.=--This change is due to the deficient formation of green +colouring matter or chlorophyll, and is more a pathological condition +than a deformity. + +It seems necessary to draw a distinction between this state and ordinary +blanching or etiolation. In the former case chlorophyll seems never to +be formed in the affected parts, even if they be exposed to light, while +an etiolated organ, when placed under favorable circumstances, speedily +assumes a green colour. In _Richardia aethiopica_ one or more leaves +become occasionally as white as the spathe is usually. + +=Virescence.=--Engelmann[373] pointed out that, so far as flowers were +concerned, there are two ways in which they assume a green colour, +either by a simple development of chlorophyll in place of the colouring +matter proper to the flower, or by an actual development of leaf-like +organs in the room of the petals--frondescence. Morren[374] judiciously +proposed to keep these two conditions separate, calling the one +virescence, the other frondescence (see p. 241). + +Many of the cases recorded as reversions of the parts of the flower to +leaves are simply instances of virescence; indeed, it is not in all +cases easy to distinguish between the two states. The examination of the +arrangement of the veins is often of assistance in determining this +point; for instance, if, under ordinary circumstances, the venation of +the petal be such as is characteristic of the sheath of the leaf, while +in the green-coloured flower of the same species the venation is more +like that which belongs to the blade of the leaf, the inference would, +of course, be that the green colour was due to frondescence or phyllody. + +The persistence or duration of petals is often increased when they are +subject to this change; instead of falling off speedily they become +persistent when so affected. + +Some flowers are more liable to virescence than others. The common +honeysuckle, _Lonicera Periclymenum_, is one of these, and it is +noticeable in this plant that the calyx remains unaffected--a +circumstance which Morren says shows the distinctness of virescence from +frondescence; for, in this instance, we have the most foliaceous portion +of the flower remaining unchanged, while the corolla and other organs, +usually less leaf-like in their nature, assume a green colour; but this +may rather be attributed to the axial nature of the so-called adherent +calyx. The stamens in these green-flowered honeysuckles are usually +green also, but with abortive anthers, and the pistil also is in a +rudimentary condition. _Umbelliferae_ are not unfrequently subject to +this change, _e.g._, _Torilis Anthriscus_, _Daucus Carota_, _Heracleum +Sphondylium_, _Carum carui_, &c. _Primulaceae_, again, are frequently +subject to virescence. Among _Compositae_ the following species are +recorded as having had green flowers--_Cirsium tricephalodes_, _Senecio +vulgaris_, _Calendula officinalis_, _Pyrethrum Parthenium_, _Carduus +crispus_, _Hypochaeris radicata_, _Hieracium prealtum_, _Cirsium +arvense_, _Coreopsis Drummondi_.[375] In _Ranunculaceae_ virescence has +been observed in _Delphinium elatum_, _crassicaule_ and _Ajacis_, +_Anemone hortensis_ and _nemorosa_, _Aquilegia vulgaris_, _Ranunculus +Philonotis_. + +Many of these cases, and others that might be cited, are probably +instances of frondescence or phyllody (see p. 241). + +=Chromatism.=--This term is here intended to apply specially to those +cases in which any organ of a plant assumes a colour approximating to +that of the petals, or in which the normal green is replaced by tints of +some other colour. To a certain extent the change in question is the +same as that spoken of under the head of petalody (see p. 283), but +there are cases in which, while the ordinary situation and form are +those of leaves, the coloration is that of the petals. Such was the case +in the _Gesnera_ mentioned by Morren (see p. 88), and in which a leaf +occupied the position of an inflorescence, and became brightly coloured. +In tulips the presence of a highly coloured leaf on the flower-stalk, +below the flower, is not uncommon. So also the bracts or leaves below +the perianth in _Anemone coronaria_ and _hortensis_ not unfrequently +assume the coloration usually confined to the parts of the perianth. A +similar illustration has presented itself, as this sheet is passing +through the press, in which two of the leaflets of the compound leaf of +a rose were brightly coloured like the petals, the others being of +their ordinary green colour. + +The occurrence of coloured bracts, as in _Poinsettia_, _Bougainvillea_, +&c., is very common under natural conditions, and need not here be +further alluded to. + +Increased intensity of colour often accompanies teratological changes; +an instance has just been alluded to in the _Gesnera_; the feather +hyacinth, _Muscari comosum_, furnishes another illustration, the +adventitious pedicels being brightly coloured. + +In fasciated stems, also, of herbaceous plants, it not unfrequently +happens that the upper portions of the stem are brightly coloured. + +The occurrence of flowers or fruits of different colours on the same +plant, or even in the same cluster, is a phenomenon which does not come +within the scope of the present book; the reader may, however, be +referred to the excellent summary on this subject published by Mr. +Darwin in his work on the 'Variation of Animals and Plants under +Domestication.' + +FOOTNOTES: + +[372] These deviations are treated of under the head of alterations of +form, because they are not, in a teratological point of view, of +sufficient importance to demand a specific heading, while they appeal to +the sight in the same way as the deviations from the customary forms of +organs. + +[373] 'De Antholys,' p. 32, Sec. 38. + +[374] 'Bull. Acad. Belg.,' xvii, part 2, p. 131, c. tab. + +[375] See Cramer, 'Bildungsabweich,' pp. 17, 55, 82, 65. See also Lucas, +'Verhandl. des Bot. Vereins. Brandenb.,' heft 1, 2, _Anchusa_. Christ, +'Flora,' 1867. pp. 376, tab. 5, 6, _Stachys_. + + + + +BOOK III. + +DEVIATIONS FROM THE ORDINARY NUMBER OF ORGANS. + + +To a certain extent the number of the organs of a plant is of even +greater consequence for purposes of classification than either their +form or their arrangement; for instance, the number of cotyledons in the +embryo is made the chief basis of separation between the two great +groups of flowering plants, the monocotyledons and the dicotyledons. In +the one group, moreover, the parts of the flower are arranged in groups +or whorls of five; in the other the arrangement is ternary. In mosses +the teeth of the peristome are arranged in fours, or in some multiple of +that number. So far as the larger groups are concerned, and also in +cases where the actual number of parts is small, the numerical relations +above described are very constant; on the other hand, in the minor +subdivisions, and especially where the absolute number of parts is +large, considerable variation may occur, so that descriptive botanists +frequently make use of the term indefinite, and apply it to cases where +the number of parts is large and variable, or, at any rate, not easy to +be estimated. + +Considered teratologically, the changes, as regards the number of +organs, are readily grouped into those consequent on a decreased and +into those resulting from an increased development. The alteration may +be absolute or relative. There may be an actual deficiency in the number +of parts or an increase in their number, but in either case the change +may be simply a restoration of the primitive number, a species of +peloria, in fact. An increased number of parts, moreover, may depend not +so much on the formation of additional parts as on the subdivision of +one. + +It seems also desirable to treat separately those cases in which there +is an increased number of buds either leaf-buds or flower-buds, as the +case may be, as happens in what is termed prolification. This formation +of buds occurring, as it does, often in unwonted situations is treated +of under the head of alterations of arrangement, the mere increase in +number being considered of subordinate importance as contrasted with the +altered disposition (see p. 100). + + + + +PART I. + +INCREASED NUMBER OF ORGANS. + + +An augmentation in the number of parts may arise from several causes, +and may sometimes be more apparent than real. True multiplication exists +simply as a result of over-development; the affected organs are repeated +sometimes over and over again each in their proper relative position, +and without any transmutation of form. + +Metamorphy, on the other hand, often gives rise to the impression that +parts are increased in number, when it may be that the stamens and +pistils, one or both, are not so much increased in number as altered in +appearance. The double anemones and ranunculus of gardens, amongst many +other analogous illustrations, may be mentioned. In these flowers, owing +to the petalody of the stamens and pistils, one or both, an impression +of exaggerated number is produced, which is by no means necessarily a +true one. Fission or lateral subdivision also gives rise to an apparent +increase in number; thus, some so-called double flowers, the elements of +which appeared to be increased in numbers, owe the appearance merely to +the laciniation or subdivision of their petals. + +The French botanists, following Dunal and Moquin, attribute an increase +in the number of whorls in the corolla, and other parts of the flower, +to a process which they call chorisis, and they consider the +augmentation to be due to the splitting of one petal, for instance, into +several;--somewhat in the same manner as one may separate successive +layers of talc one from the other. + +English botanists, on the other hand, have been slow to admit any such +process, because, in most instances, no alteration in the law of +alternation takes place in these double flowers, and in those few cases +where the law is apparently infringed, the deviation is explained by the +probable suppression of parts, which were they present would restore the +natural arrangement of the flower; and, that this is no imaginary or +purely theoretical explanation, is shown by some of the _Primulaceae_, +wherein a second row of stamens is occasionally present in the adult +condition, and renders the floral symmetry perfect. + +The double daffodil, where there are from forty to fifty petaloid organs +instead of fifteen, and wherein each piece exhibits a more or less +perfect coronal lobe at the junction of the claw and the limb, has been +cited as an objection to chorisis, though it is difficult to see on what +grounds. + +In _Delphinium_, as shown by Braun,[376] the stamens and carpels are +members of a continuous spiral series, and in the double balsam an extra +corolline whorl is produced, without the suppression of the stamens, in +the following manner: the ordinary stamens are replaced by petals, the +carpels by stamens, while an additional whorl of carpels is produced at +the summit of the axis. In this instance, therefore, the doubling is +distinctly referrible to an absolute increase in the number of whorls, +and not to chorisis.[377] + +On the other hand, it must be admitted that there are many cases which +are not to be explained in any other way than that suggested by the +French botanists before alluded to. Probably, the main difficulty in the +way of accepting the doctrine of chorisis is the unfortunate selection +of the word used to designate the process; this naturally suggests a +splitting of an organ already perfectly formed into two or more +portions, either in the same plane as the original organs, "parallel +chorisis;" or at right angles to it "collateral chorisis." Indeed, +before so much attention had been paid to the way in which the floral +organs are developed, it was thought that an actual splitting and +dilamination did really take place; Dunal and Moquin both assert as +much. The truth would rather seem to be that, in the so-called parallel +chorisis at least, the process is one of hypertrophy and +over-development rather than of splitting. The adventitious petal or +scale is an excrescence or an outgrowth from the primary organ, and +formed subsequently to it. + +In the case of "compound stamens" the original stamens are first +developed each from its own cellular "mamelon," or growing point; and, +after a time, other secondary growing points emerge from the primary +one, and in this way the stamens are increased in number, without +reference, necessarily, to the so-called law of alternation. Outgrowths +from leaves, multiplying the laminar surface, are alluded to under the +head of hypertrophy, and it is probable that some of the cases of +duplication of the flower, or of the formation of adventitious segments +outside the ordinary corolla as alluded to in succeeding paragraphs (see +Pleiotaxy of the corolla), are due to a similar process.[378] + +The formation of parts in unwonted numbers may be merely a reversion to +what is supposed to have been the original form, and in this way there +may be a restoration of parts that are usually undeveloped or +suppressed. There can be little or no doubt that there are in reality +six stamens in _Orchidaceae_, of which one only, under ordinary +circumstances, is developed. When the numerical symmetry is restored, as +it sometimes is, it is obvious that the augmentation that occurs is of a +different character from that arising from a repetition or renewed +development of organs. When the increased number arises from +multiplication proper, or from repetition, the ordinary laws of +alternation are not interfered with, but if from chorisis or +"dedoublement," it may happen that the normal arrangement is disturbed. + +Without studying the mode of development, it is not in all cases +possible to tell under which of the above categories any particular +instance should be placed; hence, in the following sections, except +where otherwise stated, the cases are grouped according to the +appearance presented in the adult condition, rather than to the way in +which the changes from the typical condition are brought about. With +reference to the foliar organs it is necessary to distinguish those +cases in which there is, from any cause, an augmentation in the number +of component parts of a whorl, from those in which the increase takes +place in the numbers of the whorls themselves. + +FOOTNOTES: + +[376] Braun, 'Pringsheim Jahrbuch f. Wiss. Bot.,' 1858, 1, p. 307, tab. +22, 23. + +[377] Henfrey, 'Jour. Linn. Soc. Bot.,' vol. iii, p. 159. + +[378] On the subject of chorisis or dedoublement the reader may +profitably consult Moquin-Tandon, 'Ess. sur les Dedoublements,' and the +same author in 'Ann. Sc. Nat.,' t. xxvii, p. 236. and 'El. Ter. Veget.,' +p. 337. Dunal, 'Consid. Org. Fleur.,' Montpell., 1829, p. 32, note 3. A. +de St. Hilaire in 'Ann. Sc. Nat.,' ser. 3, t. iii, p. 355, adnot. +Lindley, 'Elements of Botany,' p. 76. Asa Gray. 'Botanical Text Book.' + + + + +CHAPTER I. + +MULTIPLICATION OF AXILE ORGANS, INFLORESCENCE, ETC. + + +By Linne an undue number of branches was designated as "plica," from the +analogy with the disease of the hair known as plica polonica: "_Plicata +dicitur planta, cum arbor vel ramus excrescit minimis intertextis +ramulis, tanquam plica polonica ex pilis, ceu instar nidi Picae, quod +vulgo a genio ortum arbitratur; frequens apud nos in Betula, praesertim +Norlandiae, in Carpino Scaniae, nec infrequens in Pinu._"[379] + +By some of the older authors this condition was called polyclady. In +some cases, it would seem to be due to fungi as in the witches' brooms +(hexenbesen) of the German forests; in other instances, it is a result +of mutilation as after the operation of pollarding. + +Moquin-Tandon[380] mentions a case in a grafted ash in the botanic +garden of Toulouse, where below the graft there was a large swelling, +from which proceeded more than a thousand densely-packed, interlacing +branches. + +This must have been similar to the condition so commonly met with in the +birch, and frequently in the hornbeam and the thorn, and which has +prompted so many a schoolboy to climb the tree in quest of the apparent +nest. It is probable that some of the large "gnaurs" or "burrs," met +with in elms, &c., also in certain varieties of apples, are clusters of +adventitious buds, some of which might, and sometimes do, lengthen out +into branches. + +An increased number of branches also necessarily arises when the +flower-buds are replaced by leaf-buds. + +[Illustration: FIG. 179.--Flower stalks of _Bellevalia comosa_, nat. +size, after Morren.] + +Occasionally, a great increase in the number of pedicels, or +flower-stalks, may be met with in conjunction with a decreased number +of flowers, as in the wig-plant (_Rhus Cotinus_), or the +feather-hyacinth (_Bellevalia comosa_). In these cases the supernumerary +pedicels are often brightly coloured. To this condition Morren gave the +name mischomany, from [Greek: mischos], a pedicel, a term which has not +generally been adopted.[381] + +[Illustration: FIG. 180.--Tuft of branches at the end of the +inflorescence of _Bellevalia comosa_, enlarged after Morren.] + +M. Fournier[382] describes a case in the butcher's broom (_Ruscus +aculeatus_), wherein from the axil of the minute leaf subtending the +flower a secondary flattened branch proceeded. + +Duchartre[383] cites the case of a hyacinth which, in addition to the +usual scape, had a second smaller one by its side terminated by a +solitary flower; indeed, such an occurrence is not uncommon. + +Some tulips occasionally present three or four, or more, flowers on one +inflorescence, but whether from a branching of the primary scape, or +from the premature development of some of the axillary bulbils into +flowering stems which become adherent to the primary flower-stalk, +cannot, in all cases, be determined. Certainly, in some cases examined +by me the latter was the case.[384] + +Under this head, too, may be included those cases wherein an ordinarily +spicate inflorescence becomes paniculate owing to the branching of the +axis and the formation of an unwonted number of secondary buds. +Instances of this kind may be met with in willows, hazels, alders, and +other amentaceous plants. In the case of the hazel the unusual +development of male catkins sometimes coincides with an alteration in +their position, instead of being placed near the axil of a leaf; they +become terminal. Jaeger figures and describes a bunch of _Pinus +sylvestris_ bearing in one case seventy minute cones, and in another +fifty-nine. These cones preserved the same spiral arrangement among +themselves which is proper to the leaves. These latter, indeed, replaced +the strobili above.[385] + +[Illustration: FIG. 181.--Increased number of male catkins in the hazel +_Corylus avellana_.] + +M. Reichardt describes an analogous case in the same species, and +attributes the inordinate number of cones to a fungus (_Peridermium +pini_). In this case there were no less than 227 cones, but each one +half the size of the ordinary cones.[386] + +Of a similar character is the many-headed pineapple. Among grasses such +a branching of the inflorescence is exceedingly common,--which is the +more readily understood as the normal inflorescence is in so many cases +paniculate. Cultivators have, in some instances, availed themselves of +this peculiarity, as in the Egyptian wheat or corn of abundance +(_Triticum compositum_), certain varieties of Maize, etc. Similar +exuberant growths occur in _Orchidaceae_, in _Cyperaceae_, e.g. _Carex_, +in _Restiaceae_, and indeed they may be found in any plant with a similar +form of inflorescence. In all these cases the branching begins at the +lower part of the spike, and extends from below upwards in an indefinite +manner, even although the primary inflorescence be definite. + +Among the _Equisetaceae_ a similar plurality of spikes occurs often as a +result of mutilation.[387] The deviation in question might in some +instances be turned to good account, as in the _Triticum_ before +mentioned or as in the broccoli shown at fig. 182, though it must be +added that the apparent advantages are often counterpoised by some +undesirable qualities or by some circumstance which prevents us availing +ourselves of the new condition. + +=Multiplication of Bulbs.=--This occurrence has been briefly alluded to +previously (see p. 84). The most curious cases are those in which one +bulb is placed on the top of another as happened in some bulbs of +_Leucoium aestivum_ described by M. Gay.[388] Irmisch described a similar +phenomenon in _L. vernum_; and Mr. Moggridge has communicated drawings +of a similar formation in the same species grown in the neighbourhood of +Mentone. + +From the instances cited it is clear that branching of the inflorescence +occurs most frequently in those plants naturally characterised by a +dense compact mode of growth, whether that be definite or indefinite, as +in spikes, umbels, capitula, &c.; so that compound spikes, umbels, &c., +are formed in the place of simple ones (see also prolification of the +inflorescence, p. 102). + +[Illustration: FIG. 182.--Broccoli, with six perfect heads on one stalk +('Gard. Chron.,' 1856, Oct. 25).] + +=Increased number of florets= in the individual spikelets of grasses is +also met with under some circumstances. I have seen this in _Hordeum_ +and _Lolium_, and an instance is figured in _Avena_ by Dr. +Wiegmann.[389] M. Duval Jouve[390] records a similar occurrence in +_Catabrosa aquatica_, the spikelets of which contained from two to seven +flowers.[391] + +FOOTNOTES: + +[379] 'Phil Bot.,' Sec. 274. + +[380] 'El. Ter. Veget.,' p. 392. + +[381] 'Bull. Acad. Belg.,' xvii, part ii, p. 38. + +[382] 'Bull. Soc. Bot. Fr.,' vol. iv, 1857. p. 760. + +[383] Ibid., vol. viii, 1861, p. 159. + +[384] See 'Gard. Chron.,' July, 1866, p. 656, and Clusius, 'Plant. +Rar.,' lib. 2, p. 143, _Tulipa serotina_ [Greek: polyklades], _minor_, +_&c._ Hort. Eysttett. Plant. Vern.,' fol. 12. + +[385] 'Jaeger de Pini sylvestris monstrositate,' Stuttgardt, 1828. + +[386] Cited in 'Bull. Soc. Bot. Fr.,' xiv, p. 265. + +[387] Duval Jouve, 'Hist. Nat. Equiset. Fr.,' tab. 8, also Milde, 'Nov. +Act. Acad. Nat. Cur.,' t. xxvi, part 2. For branched inflorescence of +orchids, see 'Reichenbach Proc. Lond. Bot. Congress,' 1866, p. 121. + +[388] 'Bull. Soc. Bot. Fr.,' vi, 266, vii, 457. Irmisch, 'Knollen und +Zwiebelgew.,' tab. 7, figs. 10, 11. + +[389] 'Flora,' 1831, p. 5, tab. i; see also Hanstein, 'Flora,' 1857, p. +513. Schlechtendal, 'Bot. Zeit.,' xviii, p. 381. + +[390] 'Bull. Soc. Bot. Fr.,' ix, p. 8. + +[391] It will be seen, from what has been just said, that in some of the +cases where the axile organs, branches, &c., appear to be multiplied, +the increased number is due to subdivision rather than to renewed +formation (see Fission). Of this last description is an instance which +came under the writer's notice after the section relating to that +subject was in print, and which may therefore here be alluded to. The +instance is that of the subdivision of the leaf-like organs of +_Sciadopitys verticillata_. In one instance the pseudo leaf divided, and +from the division proceeded a little axis, bearing at its summit a +verticil of pseudo leaves. This division and formation of new axes and +verticils affords ample confirmation of the opinion thrown out by +Professor Alexander Dickson, that the apparent leaves of this plant were +really branches: see 'Revue Horticole,' 1867, and 'Report. Bot. +Congress,' London, 1866, p. 124. + + + + +CHAPTER II. + +MULTIPLICATION OF FOLIAR ORGANS. + + +The cases referrible to this head may be ranged under two sections +according as the increase is due to plurality of ordinarily single +organs, or to an increase in the number of verticils or whorls. + +When, in place of a single leaf organ two or more are really or in +appearance present the occurrence may be due to one of several causes; +among them may be mentioned an actual formation of parts in unwonted +number, hypertrophy or enation, chorisis or fission, disjunction, +adhesion of one leaf to another or to the stem, as in some of the leaves +called "geminate," wherein the two leaves, though apparently in +juxtaposition, yet originate from different parts of the stem, but by +coalescence or lack of separation produce the impression as if they +sprang from the same node. In the adult state it is not always possible +to ascertain with certainty to which of these causes the increase in the +number of leaves is due, though a clue to the real state of things may +be gained from attention to the distribution of the veins, to the +arrangement or phyllotaxy of the leaves, the size and position of the +supernumerary organs, &c. + +The term "phyllomania," as ordinarily used, is applied to an unwonted +development of leafy tissue, as in some begonias where the scales or +ramenta are replaced by small leaflets, or as in some cabbage leaves, +from the surface of which project, at right angles to the primary +plane, other secondary leafy plates; but these are, strictly speaking, +cases of hypertrophy (see Hypertrophy). + +Those instances in which the actual number of leaves is increased, so +that in place of one there are more leaflets, may be included under the +term "pleiophylly," which may serve to designate both the appearance of +two or more leaves in the place usually occupied by a single one, and +also those normally compound leaves in which the number of leaflets is +greater than usual. + +The increased number of leaves in a whorl may well be designated as +"polyphylly," using the word in the same sense as in ordinary +descriptive botany, while "pleiotaxy" may be applied to those cases in +which the number of whorls is increased. + +[Illustration: FIG. 183.--Supernumerary leaflet, _Ulmus campestris_.] + +=Pleiophylly.=--As above stated, this term is proposed to designate +those cases in which there is an absolute increase in the number of +leaves starting from one particular point, as well as those in which the +number of leaflets in a compound leaf is preternaturally increased. The +simplest cases are such as are figured in the adjacent cuts, wherein, in +place of a single leaf, two are produced in the elm. In the one case the +new leaflet springs from the apex of the petiole and partially fills +the space consequent on the obliquity of the base of the leaf. In the +other it would seem as if two distinct leaves emerged from the stem in +juxtaposition. This is probably due to a lateral chorisis or subdivision +of the primitive tubercle or growing point, followed by a like +subdivision of the vascular bundle supplying it. There are certain +varieties of elm that very generally present this anomaly on their rank, +coarse, growing shoots. In these cases the new growths have the same +direction as the primary one, but in other cases the supplementary +production is exactly reversed in direction. Thus, in the common hazel +(_Corylus_) a second smaller leaf proceeding from the end of the +leaf-stalk at the base of the primary one may frequently be seen. M. +Germain de Saint Pierre records an instance in a mulberry leaf, from the +base of which proceeded a large leafy expansion divided into two +tubular, horn-like projections, and in the centre a thread-like process +representing the midrib and terminated by a small two-lipped limb.[392] +Dr. Ferdinand Mueller speaks of a leaf of _Pomaderris elliptica_ as +bearing a secondary leaf on its under surface.[393] + +[Illustration: FIG. 184.--Supernumerary leaf, _Ulmus montana_.] + +[Illustration: FIG. 185.--Supernumerary leaf of hazel.] + +The leaves of _Heterocentron macrodon_ have likewise been observed +occasionally to produce leaflets from their upper surface. + +To this production of leaves from leaves the late Professor Morren +applied the term "autophyllogeny."[394] The Belgian botanist figures a +small perfect leaf springing from the nerves of the upper surface of the +primary leaf in a species of _Miconia_. As in the hazel, the direction +of the adventitious leaf is inversely that of the primary one, the upper +surface of the supernumerary leaflet being turned towards the +corresponding surface of the normal leaf. A similar occurrence took +place in _Gesnera zebrina_, but the new growth in this case sprang from +the lower face of the leaf. Morren explains the appearances in question +by supposing that the supplementary leaf is one of a pair belonging to a +bud borne on a slender stalk. This stalk and one of the bud-leaves are +supposed to be inseparably united with the primary leaf. But there is +no reason at all for supposing the existence of adhesion in these cases; +no trace of any such union is to be seen. A much more natural +explanation is that, from some cause or another, development at the apex +of the petiole or on the surface of the nerves, instead of taking place +in one plane only, as usual, takes place in more than one, thus showing +the close relationship, if not the intrinsic identity, between the +leaf-stalk and its continuation, the midrib, with the branch and its +subdivisions. The form of the leaf-stalk and the arrangement of the +vascular bundles in a circle in the case of the hazel, before alluded +to, bear out this notion. Such cases are significant in reference to the +notion propounded by M. Casimir de Candolle, that the leaf is the +equivalent of a branch in which the upper portion of the vascular circle +is abortive.[395] + +Compound leaves, as has been stated, occasionally produce an extra +number of leaflets; one of the most familiar illustrations of this is in +the case of the four-leaved shamrock (_Trifolium repens_), which was +gathered at night-time during the full moon by sorceresses, who mixed it +with vervain and other ingredients, while young girls in search of a +token of perfect happiness made quest of the plant by day. Linne, who in +this matter, at any rate, had less than his usual feeling for romance, +says of the four-leaved trefoil that it differs no more from the +ordinary trefoil than a man with six fingers differs from one provided +with the ordinary number. It should be stated that five and six +adventitious leaflets are found almost as frequently as four. + +Walpers describes a case where the leaf of _T. repens_ bore seven +leaflets. Schlechtendal alludes to a similar increase in number in +_Cytisus Laburnum_, and many other instances might be cited. + + For figures or descriptions of four-leaved shamrocks the reader + is referred to Lobel, 'Stirp. Advers.,' Nov., p. 382. + Tabernaemontanus 'Krauterbuch,' S. 222. Schlechtendal, 'Bot. + Zeit.,' ix, p. 583, xiv, p. 71. Maugin, 'Bull. Soc Bot. Fr.,' + 1866, t. xiii, p. 279. See also Cramer, 'Bildungsabweich,' p. + 92. Walpers, 'Linnaea,' 1840, p. 362 (7-leaved). Schlechtendal, + 'Bot. Zeit.,' 1844, p. 457, _Cytisus_. Wigand, 'Flora,' 1856, + p. 706. + +Frondiferous leaves have much the appearance of branches provided with +leaves, and they may be compared with those instances in which an +adventitious bud is placed on the surface or edges of the leaves, as in +_Gesnera_, _Cardamine_, &c. In truth, the two conditions merge one into +the other, as in some begonias, where the ramenta often become leaf-like +and bear small bulbils in the axil. + +When frondiferous leaves die the appendages die also, but when a true +bud has been formed on a leaf it does not of necessity die with the leaf +that bears it, but separates from it and continues to grow +independently. + +=Increased number of stipules, spathes, &c.=--Seringe relates the +occasional presence of two or three additional stipules upon the +leaf-stalks of _Salix fragilis_, and even makes a variety (_Salix +pendula_, var. _multistipulata_). + +An increase in the number of the spathes has been often noticed in +Arads[396]. Prof. Alex. Braun has studied this subject in some +detail[397]. In _Calla palustris_ the shoot which continues the growth +of the plant proceeds from the axil of the last leaf but one; the very +last leaf producing no bud, but if accidentally a shoot is developed in +this latter situation it produces flowers at once. No leaves are formed, +but, on the contrary, two or three spathes surround the spadix, so that +the presence of an increased number of spathes in this plant is +associated with the development of a side shoot from the axil of the +last leaf, the situation whence, under natural circumstances, no shoot +at all issues. The supernumerary spathes are not always on the same +level, but may be separated by a considerable interval. They vary very +much in size, and sometimes assume the form and appearance of leaves. +Similar anomalies occur in other Arads as _Arum maculatum_, _Richardia +aethiopica_, and _Anthurium Scherzerianum_, frequently combined with a +leaf-like appearance of the spathes and sometimes with a subdivision of +the spadix into two or three branches. + +Engelmann relates the occurrence of an increased number of glumes in +_Bromus velutinus_ associated with suppression of the flowers. + +=Polyphylly.=--As previously explained, this term is here applied to +those cases in which the members of any particular whorl are increased +in number, the whorls themselves not necessarily being augmented. + +The simplest cases of this kind are those in which we meet with an +unusual number of leaves in a whorl. + +=Increased number of leaves in a whorl.=--This may arise from actual +multiplication, or from lateral chorisis, or fission. The true nature of +the case may usually be ascertained by an examination of the +distribution of the veins of the leaves, or of the fibrous cords of the +stem, by the relative position of the supernumerary organs, &c. + +Among plants with normally opposite leaves the following occasionally +produce them in whorls of three:--_Lonicera brachypoda_, _L. Xylosteum_, +_Weigela rosea_, _Cornus mas_, _Vinca minor_, &c. + +_Paris quadrifolia_ may frequently be met with five leaves in its whorl, +or even six.[398] + +=Increased number of bracts.=--This is not of infrequent occurrence; one +of the most curious instances is that recorded by Mr. Edwards[399] in +_Cerastium glomeratum_, where, in place of the usual pair of bracts at +the base of the head of flowers, there was a whorl of six or eight, +forming an involucre. The flowers in this case were apetalous and +imperfect. + +=Polyphylly of the calyx.=--This may occur without any other +perceptible change, while at other times the number of the other parts +of the flower is proportionately increased. In a flower of a plum six +sepals in place of five sometimes exist; a precisely similar occurrence +in the flowers of the elder (_Sambucus_), the _Fuchsia_, and of +_OEnanthe crocata_, may occasionally be met with. In the latter case, +indeed, there are sometimes as many as ten segments to the calyx, and +this without the other parts of the flower being correspondingly +augmented. Among monocotyledons a similar increase is not uncommon, as +in _Tulipa_, _Allium_, _Iris_, _Narcissus_, &c. + +In some plants there seems to exist normally much variation in the +number of parts; thus in some species of _Lacistema_ in adjacent flowers +the calyx may be found with four, five, or six segments. + +Most of these cases of polyphylly affecting the calyx may be explained +by lateral chorisis or fission. + +=Polyphylly of the corolla.=--This may happen in connection with similar +alterations in the calyx and stamens, or sometimes as an isolated +occurrence. In the latter case it may be due to lateral chorisis, to +substitution, or to the development of organs usually suppressed; thus, +when in aconites we meet with four or five horn-like nectaries (petals) +instead of two only, as usual, the supernumerary ones are accounted for +by the inordinate development of parts which ordinarily are in an +abortive or rudimentary state only. This is borne out by what happens in +_Balsamineae_. In the common garden balsam the fifth petal is +occasionally present, while in _Hydrocera triflora_ this petal is always +present. + +In a flower of a _Cyclamen_ recently examined there were ten petals in +one series, the additional five being evidently due to the subdivision +of the five primary ones; the natural circular plan of the flower was +here replaced by an elliptical one. A similar occurrence takes place in +the flowers of maples (_Acer_), which sometimes show an increased number +of parts in their floral whorls and an elliptical outline. Whether the +additional organs in this last case are the result of complete lateral +chorisis or of multiplication proper I do not know. + +Orchids are very subject to an increase in the number of their labella. +As illustrations may be cited an instance recorded by Mr. J. T. +Moggridge in a flower of _Ophrys insectifera_, and in which there were +two labella without any other visible deviation from the ordinary +conformation.[400] + +I am indebted to Mr. Hemsley for the communication of a similar specimen +in _O. apifera_, in which there were two divergent lips, each with the +same peculiar markings. One of the sepals in this flower was adherent to +one of the lateral petals. This augmentation of the labella depends +sometimes on the separation, one from the other, of the elements of +which the lip is composed, at other times on the development, in the +guise of lips, of stamens which are usually suppressed (see p. 380). + +The following enumeration will suffice to show the genera in which an +increased number of petals or perianth-segments in any given whorl most +frequently occurs. + + Anemone! + Ranunculus! + Aconitum! + Raphanus. + Bunias. + Saponaria. + Dianthus! + Pelargonium! + Hibiscus. + Fuchsia. + Sarothamnus! + Lotus! + Ulex! + Prunus! + Trifolium. + OEnanthe and Umbellif. pl.! + Sambucus! + Bryonia. + Campanula. + Solanum. + Veronica. + Cyclamen! + Primula! + Anagallis! + Plumbago. + Jasminum. + Syringa! + Tradescantia. + Iris. + Tigridia. + Narcissus. + Tulipa. + Convallaria! + Paris! + Hyacinthus! + Allium! + Ornithogalum. + Orchideae, sp. pl.! + + For other illustrations see multiplication of whorls, petalody; + see also Moquin, loc. cit., p. 350. Engelmann, loc. cit., p. + 20, Sec. 18. Cramer, loc. cit., p. 25. + +=Polyphylly of the androecium.=--An increased number of stamens +frequently accompanies the corresponding alterations in other whorls, +and seems, if anything, to be more frequent among monocotyledonous +plants than among dicotyledonous ones; thus, we occasionally find +tetramerous flowers in _Crocus_, _Hyacinthus_, _Tulipa_, _Iris_, +_Tigridia_, &c., and more rarely in _Yucca_ (_Y. flexilis_[401]). + +The increased number of stamens in a single whorl may result from a +development of organs usually suppressed, and constitute a form of +regular peloria as in _Linaria_, wherein a fifth stamen is occasionally +met with. Among normally didynamous plants such numerical restitution, +so to speak, is not unusual; thus, in _Veronica_ four and five stamens +occur. Fresenius has seen five stamens in _Lamium_, _Mentha_, +_Chelone_;[402] Bentham in _Melittis_, and other instances are cited +under the head of peloria. Chorisis may also serve to account for some +of these cases; thus, Eichler[403] figures a flower of _Matthiola annua_ +with five long stamens instead of four; one of the long pairs of stamens +has here undergone a greater degree of repetition than usual. De +Candolle[404] cites and figures a curious form of _Capsella +Bursa-pastoris_ sent him by Jacquin, and which was to some extent +reproduced by seed. In the flowers of this variety there were no petals, +but ten stamens; hence De Candolle inferred that the petals were here +replaced by stamens, but Moquin[405] objects, and with justice, to this +view, as the ten stamens are all on the same line; he considers the +additional stamens to be the result of chorisis. Buchenau[406] mentions +the presence of seven stamens in another Crucifer, _Ionopsidium +acaule_. Here the supernumerary organ was placed between two of the long +stamens. The effect of chorisis in producing an augmentation of parts is +well seen in some plants that have some of their flowers provided with +staminodes or abortive stamens, and others with clusters or phalanges of +perfect stamens. Thus, in the female flowers of _Liquidambar_ there are +five small staminodes without anthers, whereas in the male flower the +stamens are numerous and grouped together in phalanges, so that the +relation of simple to compound stamens is in this case readily seen, as +also in many _Malvaceae_, _Sterculiaceae_, _Byttneriaceae_, _Tiliaceae_, and +_Myrtaceae_. It is probably the idea of splitting or dilamination +involved in the word chorisis that has led many English botanists to +hesitate about accepting the notion. Had they looked upon the process as +identical with that by which a branched inflorescence replaces an +unbranched one, or a compound leaf takes the place of a simple one, the +objections would not have been raised with such force. The process +consists, in most cases, not so much in actual cleavage of a +pre-existing organ as in the development of new-growing points from the +old ones. + +An illustration given by Moquin from Dunal[407] goes far to support the +notion here adopted. The majority of the stamens of laurels (_Laurus_) +have, says M. Dunal, on each side of the base of their filaments a small +glandular bifid appendage; these excrescences are liable to be changed +into small stamens. The male flowers have a four-leaved calyx, and +sometimes eight stamens, each with two glands, four in one row, opposite +to the sepals, four in a second series alternating with the first. More +generally two of the stamens are destitute of glands, but have in their +place a perfectly developed stamen, so that in these latter flowers +there are twelve stamens. + +M. Clos[408] mentions a flower of rue (_Ruta_) wherein there were two +stamens joined together below and placed in front of a petal, as in +_Peganum_. + +Buchenau[409] mentions a flower of _Lotus uliginosus_ in which there +were eleven stamens, namely, two free and nine monadelphous; and +Hildebrand describes an analogous increase in a flower of _Sarothamnus +scoparius_ in which, in conjunction with a seven-toothed calyx, there +were two carinas and fourteen stamens. It would seem probable in this +case that there was a coalescence of two flowers at an early date and +consequent suppression of some of the parts of the flower. Whether this +was the case or not in this particular illustration, it is nevertheless +certain that many of the recorded instances of increased number in the +organs of a flower are really the results of a fusion of two or more +flowers, though frequently in the adult state but few traces of the +coalescence are to be seen. + +=Polyphylly of the gynoecium.=--Moquin[410] remarks that, as the +pistils are, generally speaking, more or less subject to pressure, owing +to their central position, and it may be added owing to their later +development, than the other parts of the flower, they are more subject +to suppression than to multiplication; nevertheless, augmentation in the +number of carpels does occasionally take place, especially when the +other parts of the flower are also augmented in number. Sometimes this +increase in the number of carpels is due to pure multiplication, without +any other change. At other times the increase is due to a substitution +of stamens or other organs for carpels (see Substitutions). In other +cases the augmentation seems to be due to the development of parts +usually suppressed; for instance, in _Antirrhinum_, where there are +usually only two carpels present, but where, under peculiar +circumstances, five may be found--thus rendering the symmetry +complete.[411] In _Papilionaceae_, wherein usually only one carpel is +developed, we occasionally find two, or even more, as in _Wistaria_, +_Gleditschia_, _Trifolium_, &c. In _Prunus_ and _Amygdalus_ from two to +five carpels are occasionally to be found,[412] in _Mimosa_ five, in +_Umbelliferae_ three to five; in some composites, _e.g._ _Spilanthes_, +five carpels have also been noticed; in _Cruciferae_ three and four, in +grasses three.[413] The double cocoa-nut affords an illustration of the +development of two carpels out of three, one only generally arriving at +perfection. Triple nuts (_Corylus_) also owe their peculiarity to the +equal development of all three carpels which exist in the original +flower, but of which, under ordinary circumstances, two become abortive. +It is necessary, however, to distinguish these cases from those in which +two embryos are developed in one seed. + +The following list may serve to show in what genera this change has been +most frequently noticed, and it may be said in general terms that +_Cruciferae_, _Umbelliferae_, and _Liliaceae_, are the orders most +frequently affected. Cases of peloria are not included in the subjoined +list. + + Nigella. + Aquilegia. + Paeonia! + Delphinium! + Iberis. + Diplotaxis. + Lunaria. + Ricotiana. + Octadenia. + Draba! + Lepidium. + *Cheiranthus! + Dianthus. + Brassica! + Parnassia. + *Acer! + Ptelea. + Citrus! + Philadelphus. + Prunus! + Amygdalus! + Crataegus! + Fuchsia! + Trapa! + Cassia. + Cercis. + Medicago. + *Phaseolus! + Wistaria. + Gleditschia. + Affonsea. + Trifolium! + Archidendron. + Mimosa. + Robinia. + Diphaca. + Coesalpinia. + Vicia. + Anthyllis. + Cucurbita. + Passiflora! + Sambucus! + *OEnanthe! + Daucus! + Angelica! + Heracleum! + Silaus. + Carum. + Thysselinum. + Campanula! + Spilanthes. + Chrysanthemum. + Anagallis. + Primula! + Fraxinus! + Lycium. + Cobaea. + Datura! + Solanum! + Sesamum. + Sideritis. + Coleus. + Veronica! + *Digitalis! + Antirrhinum! + Linaria. + Gloxinia! + Symphytum. + Anchusa. + Polygonum. + Euphorbia. + Cneorum. + Mercurialis! + Chenopodium. + Suaeda. + Beta. + Corylus! + Lambertia. + Cocos! + Tigridia. + Tulipa! + Iris! + Narcissus! + Allium! + Ornithogalum. + Gagea! + Tradescantia! + Schoenodon. + Bambuseae. + + A few additional references may here be given to papers where + an increased number of carpels is described:--Engelmann, 'De + Antholys,' Sec. 17, p. 19. Bernhardi, 'Flora,' 1838, p. 129. + Schkuhr., 'Bot. Handb.,' t. 179. Godron, 'Ann. Sc. Nat.,' ser. + 5, vol. ii, p. 280, tab. xviii, _pluricarpellary Crucifers_. + Weber, 'Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,' &c., + 1860, _Cerasus_, &c., &c. Baillon, 'Adansonia,' iv, p. 71, + _Trifolium_. Schlechtendal, 'Bot. Zeit.,' xv, p. 67, _Datura_, + three-celled fruit; 'Bot. Zeit.,' xiii, p. 823, _Phaseolus_, + double pistil--a common case. Cramer, 'Bildungsabweich,' p. 99, + reference to several leguminous plants with polycarpellary + pistils. Munro, Gen., 'Linn. Trans.,' vol. xxvi, p. 26, + _Bambuseae_. Alph. de Candolle, 'Neue Denkschrift,' + _Cheiranthus_. Schimper, 'Flora,' 1829, ii, p. 433. Wigand, + 'Bot. Untersuch.' Fleischer, 'Missbild. Cultur Pfl.' Cramer, + 'Bildungsabweich,' p. 65, _Umbelliferae_. + +=Polyphylly of the flower in general.=--Although, for the sake of +convenience, multiplication has here been treated of as it affects the +members of individual whorls of the flower, yet it must be remembered +that, in general, the augmentation is not confined to one whorl, but +affects several; thus, if the sepals are increased, the petals are +likely to be so likewise, and so forth. One of the most curious +illustrations of this is that recorded by Mr. Berkeley[414] in a plum, +wherein there was an increased number of sepals, a corresponding +augmentation in the petals, while the pistil was composed of two and +sometimes three carpels distinct from the calyx and from each other. In +the flowers there did not appear to be any definite relation in the +position of the parts either with reference one to another or to the +axis. + +[Illustration: FIG. 186.--Plum. Increased number of parts in the +calycine, corolline, and carpellary whorls respectively.] + +In _Primulaceae_ this general augmentation has been frequently +noticed.[415] + +Among _Orchideae_ the instance related by Dr. Seubert is worth alluding +to here. This botanist observed and figured a flower of _Orchis +palustris_ with tetramerous arrangement of parts, that is to say there +were four outer segments to the perianth, four petals, of which two +were lip-like, four stamens, three of which were rudimentary, and an +ovary with four parietal placentae.[416] + +The following list will serve to show in what plants this general +augmentation of parts has been observed most frequently: + + Ranunculus. + Clematis! + Delphinium. + Brassica! + Ruta. + Acer! + Prunus! + Rosa! + Rubus. + Philadelphus! + Chrysosplenium. + Umbelliferae, sp. pl.! + *Fuchsia! + OEnothera. + Adoxa. + Bryonia. + Cucumis! + Campanula! + Sambucus! + *Primula! + Anagallis! + Lycium. + Solanum. + Symphytum. + Syringa! + Linaria. + Chenopodium. + *Paris! + Convallaria! + Allium. + *Lilium! + *Tulipa! + Ornithogalum. + *Gagea! + Tradescantia! + Orchideae, sp. pl.! + +=Increased number of ovules or seeds.=--This appears not to be of very +frequent occurrence, at least in those plants where the number of these +organs is normally small; where, as in _Primula_, the ovules and seeds +are produced in large quantities, it is not practicable to ascertain +whether the number be augmented or not in any particular case. Very +probably, the attachment or source of origin of the ovules determines, +in some measure, their number. Thus, in the case of marginal +placentation the number must be limited by the narrow space from which +they proceed, whereas in parietal and free central placentation the +ovules are generally numerous. In the latter case, however, it will be +remembered that solitary ovules are not rare. An increased number of +ovules is generally remarked in conjunction with some other change, such +as a foliaceous condition of the carpel, in which the margins are +disunited. In such cases the ovules may occupy the margin or may be +placed a short distance within it, as in the case of some open carpels +of _Ranunculus Ficaria_,[417] and in which two ovules were borne in +shallow depressions on the upper or inner surface of the open carpel and +supplied with vascular cords from the central bundle or midrib. The +outer coating of the ovule here contained barred or spiral fusiform +vessels derived from the source just indicated. + +In the very common cases where the pistil of _Trifolium repens_ becomes +foliaceous (see Frondescence), the outer ovules are generally two or +more instead of being solitary. So, also, in the Rose with polliniferous +ovules (see p. 274). Among _Umbelliferae_ affected with frondescence of +the pistil a similar increase in the number of ovules takes place. It +will be borne in mind that in most, if not all, these cases the +structure of the ovule is itself imperfect.[418] + +What are called in popular parlance double almonds or double nuts +(_Corylus_) are cases where two seeds are developed in place of one. + +In the 'Revue Horticole,' 1867, p. 382, mention is made of a bush which +produces these double nuts each year--in fact, it never produces any +single-seeded fruit. The plant was a chance seedling, perhaps itself the +offspring of a double-seeded parent. It would be interesting to observe +if the character be retained by the original plant, and whether it can +be perpetuated by seed or by grafting. + +It is necessary to distinguish in the case of the nut between additional +seeds or ovules, as just described, and the double, triple, or fourfold +nuts that are occasionally met with, and which are the result either of +actual multiplication of the carpels or of the continued development of +some of the carpels which, under ordinary circumstances cease to grow +(see _ante_, p. 364). In the case of a ripe nut with two seeds it might +be impossible to tell whether the adventitious seed were the product of +multiplication, or whether it belonged, in the first instance, to the +same carpel as that producing the fellow-seed, or to a different and now +obliterated ovary. In all probability, however, the second seed would be +accounted for by the development of two seeds in one carpellary cavity. + +There is still another condition occasionally met with in the almond, +and which must be discriminated from the more common multiplication of +the seed, and which is the multiplication of the embryos within the +seed, and which furnishes the subject of the succeeding paragraph. + +=Increased number of embryos.=--A ripe seed usually contains but a +single embryo, although in the ovular state preparation is commonly made +for more; and, indeed, in certain natural orders plurality of embryos in +the same seed does occur, as in _Cycadeae_ and _Coniferae_. In the seeds +of the orange (_Citrus_), in those of some _Euphorbiaceae_, &c., there +are frequently two or more additional embryos. A similar occurrence has +been recorded in the mango, for a specimen of which I am indebted to the +Rev. Mr. Parish, of Moulmein.[419] + +Plurality of embryos has also been observed in-- + + Raphanus sativus. + *Citrus Aurantium! + Diosma, sp. + Hypericum perforatum. + Triphasia aurantiaca. + *AEsculus Hippocastanum! + Euonymus latifolius. + *Mangifera indica! + Eugenia Jambos. + Amygdalus vulgaris! + Vicia, sp. + Cassia, sp. + *Viscum album! + Daucus Carota. + Ardisia serrulata! + Cynanchum nigrum. + fuscatum. + Euphorbia rosea. + Coelebogyne ilicifolia. + Allium fragrans. + Funckia, sp. + Carex maritima. + Zea Mays. + + See Schauer's translation of Moquin-Tandon, 'El. Terat. + Veget.,' p. 245, adnot., and 'Al. Braun Polyembryonie.' + +=Increased number of the cotyledons.=--Although the presence of one or +of two cotyledons in the embryo is generally accepted as a valuable +means of separating flowering plants into two primary groups, yet, like +all other means of discrimination, it occasionally fails, and, indeed, +almost always requires to be taken in conjunction with some other +character. There are cases among flowering plants where the embryo is +homogeneous in its structure, there are others in which the number of +the cotyledons is more than two. Thus, in some seeds of _Cola acuminata_ +the cotyledons vary in number from two to five. I have not been able to +ascertain precisely whether this multiplication of the cotyledons is +characteristic of all the seeds of particular trees, or whether some +only are thus affected. Some fruits that I examined bore out the latter +view, as in the same pod were seeds with two, three, and four cotyledons +respectively. + +I have also seen three cotyledons present in embryo-plants of _Correa_, +_Crataegus Oxyacantha_, _Dianthus sinensis_, _Daucus Carota_, _Cerasus +Lauro-cerasus_. De Candolle alludes to a case of the kind in the bean, +and figures a species of _Solanum_ with three cotyledons.[420] Jaeger +alludes to a similar instance in _Apium Petroselinum_;[421] Ehrenberg to +one in the marigold (_Calendula_);[422] Reinsch to an analogous +appearance in the beech (_Fagus_), associated with a union of the +margins of two out of the three cotyledons, and of those of two out of +the three leaves next adjacent.[423] This fusion seems frequently to +accompany increase in the number of cotyledons. It was so in the +_Correa_, and in the _Crataegus_ previously mentioned. Some of these +cases may be accounted for by chorisis or by a cleavage of the original +cotyledons, as happens, according to Duchartre,[424] in some Coniferae, +which he considers to be improperly termed polycotyledonous. Whether +this holds good in the Loranths, where (_Nuytsia_, _Psittacanthus_) an +appearance of polycotyledony exists, is not stated. In the case of the +rue (_Ruta_) figured by M. A. de Jussieu[425] this splitting of one +cotyledon into two is sufficiently evident, as is also the case in the +sycamore (_Acer pseudo-platanus_), seedlings of which may often be met +with divided cotyledons. + +In other instances a fusion of two embryo plants may give rise to a +similar appearance, as in the _Euphorbia_ and _Sinapis_ found by M. +Alph. de Candolle (see _ante_, p. 56). + +=Pleiotaxy or multiplication of whorls.=--In the preceding section +notice has been taken of the increased number of parts in a single +whorl, but an augmentation of the number of distinct whorls is still +more frequently met with. Many of the so-called double flowers owe their +peculiarity to this condition. The distinction between the two modes in +which the parts of the flower are increased in number has been pointed +out by Engelmann, Moquin, and others, and the two seem to require +distinctive epithets; hence the application of the terms polyphylly and +pleiotaxy, as here proposed. + +=Pleiotaxy in the bracts.=--An increase in the number of bracts has been +met with very constantly in a species of _Maesa_, and in a peculiar +variety of carnation, called the wheat-ear carnation.[426] In some of +these cases the increase in the number of bracts is attended by a +corresponding suppression in the other parts of the flower. Such a +condition has been frequently met with in _Gentiana Amarella_, where the +bracts are increased in number, coloured purple, and destitute of any +true floral organs. A similar condition exists in some varieties of +_Plantago major_ (var. _paniculata_), as has been previously stated, p. +109. + +[Illustration: FIG. 187.--Wheat-ear carnation. The appearance is due to +the multiplication of the bracts and the suppression of the other parts +of the flower.] + +It has been noticed also in the common pea, _Pisum sativum_, and M. +Lortet[427] records a case of the kind in _Erica multiflora_, the +flowers of which, under ordinary circumstances, are arranged in +clusters, but in this case the pedicels were more closely crowded than +usual, and were covered for their whole length with small rose-coloured +bracts arranged in irregular whorls, the upper ones sometimes enclosing +imperfect flowers. In the 'Gardeners' Chronicle,' 1865, p. 769, is +figured a corresponding instance of _Delphinium Consolida_, in which the +bracts were greatly increased in number, petaloid, and, at the same +time, the central organs of the flower were wholly wanting. + +[Illustration: FIG. 188.--_Delphinium Consolida_. Multiplication of +bracts at the expense of the other parts of the flower.] + +[Illustration: FIG. 189.--Multiplication of bracts, &c., _Pelargonium_.] + +In flowers of _Pelargonium_ may occasionally be seen a repetition of the +whorls of bracts, in conjunction with suppression and diminished size of +some of the other portions of the flower (fig. 189). + +The common foxglove (_Digitalis purpurea_) has likewise occasionally +been observed subject to a similar malformation. + +_Cornus mas_ and _C. suecica_ sometimes show a triple involucre.[428] +Irmish[429] records an analogous case in _Anemone Hepatica_, wherein the +involucre was doubled. Similar augmentation occurs in cultivated +Anemone. In addition to the plants already mentioned, Engelmann[430] +mentions as having produced bracts in unwonted numbers, _Lythrum +Salicaria_, _Plantago major_, _Veronica spicata_, _Echium vulgare_, +_Melilotus arvensis_, and _Rubus fruticosus_. + +It must here be remarked that this great number of the bracts occurs +naturally in such plants as _Godoya_, in which the bracts, or, as some +consider them, the segments of the calyx, are very numerous, and +arranged in several overlapping segments. + +In some of the cultivated double varieties of _Nigella_ the finely +divided involucral bracts are repeated over and over again, but on a +diminished scale, to the exclusion of all the other parts of the flower. + +=Pleiotaxy or repetition of the calyx.=--The true calyx is very seldom +affected in this manner, unless such organs as the epicalyx of mallows, +_Potentilla_, &c., be considered as really parts of the calyx. + +In _Linaria vulgaris_ Roeper observed a calyx consisting of a double +series, each of five sepals, in conjunction with other changes.[431] It +is also common in double columbines, delphiniums, nigellas, &c. + +In the 'Revue Horticole,' 1867, p. 71, fig. 9, is described and figured +by M. B. Verlot a curious variety of vine grown for years in the Botanic +Garden at Grenoble, under the name of the double-flowered vine. The +place of the flower is occupied by a large number of successive whorls +of sepals disposed in regular order, and without any trace of the other +portions of the flower. It is, in fact, more like a leaf-bud than a +flower. The outermost whorls of this flower open at the time when the +ordinary flowers of vines do; the second series are gradually produced, +and expand about the time when the ovaries of the normal flowers begin +to swell; a third series then gradually forms, and so on, until frost +puts a stop to the growth. This malformation, it appears, is produced +annually in certain varieties of vine, and may be perpetuated by +cuttings. + +The flower of the St. Valery apple, already alluded to under the head of +sepalody, might equally well be placed here. It is not very material +whether the second whorl of organs be regarded as a repetition of the +calyx or as a row of petals in the guise of sepals. + +Engelmann[432] cites the following plants as occasionally presenting a +repetition of the calyx, in most cases with a suppression of the other +floral whorls:--_Stachys lanata_, _Myosotis palustris_, _Veronica +media_, _Aquilegia vulgaris_, _Nigella damascena_, _Campanula +rapunculoides_. + +=Pleiotaxy in the perianth.=--Increase in the number of whorls in the +perianth is common in lilies, narcissus, hyacinths, &c. It may be also +met with occasionally among orchids. The lily of the valley +(_Convallaria maialis_) seems also to be particularly subject to an +increase in the number of parts of which its perianth consists, the +augmentation being due partly to repetition or pleiotaxy, partly to the +substitution of petaloid segments for stamens and pistils.[433] + +In this place may also be mentioned the curious deviation from the +ordinary structure occasionally met with in _Lilium candidum_, and known +in English gardens as the double white lily. In this case there are no +true flowers, but a large number of petal-like segments disposed in an +irregular spiral manner at the extremity of the stem, some of the +uppermost being occasionally verticillate.[434] + +[Illustration: FIG. 190.--Double white lily. Multiplication of +perianth-segments and other changes.] + +=Pleiotaxy of the corolla.=--With reference to double flowers, it was +remarked by Linne that polypetalous flowers were, as he said, +multiplied, while monopetalous flowers were duplicated, or triplicated, +as the case may be,[435] a statement that is true in the main, though +it requires modification. In the case of polypetalous, or rather +dialypetalous flowers, the petals may be very largely increased by +multiplication, as in roses, anemones, pinks, &c. In the last-named +genus the number is often so much increased that the calyx splits from +the tension exercised on it by the increasing mass within. This +multiplication may happen without any metamorphy or substitution of +petals for stamens, though, in the majority of cases, it is associated +with such a change. It is curious to observe in some of these flowers +that the total number of parts is not greatly increased; thus, in some +of the double-flowered _Leguminosae_, such as _Ulex europaeus_ and _Lotus +corniculatus_, the petals are repeated once or twice, the stamens are +petalodic, but reduced in number, while the carpels are usually entirely +wanting. Thus, owing to the diminished number of parts in the inner +whorls of the flower, these very double-looking blooms do not contain +any greatly increased number of parts.[436] + +Flowers that, under ordinary circumstances, are gamopetalous, become, in +some instances, multiplied by the formation of additional segments, just +as in the case of polypetalous corollas; but in these cases the corollas +become polypetalous, their petals do not cohere one with another. Among +double flowers of this character may be mentioned _Campanula +rotundifolia_, _Gardenia_ sp., _Nerium Oleander_, _Serissa_ sp., +_Arbutus Unedo_, &c. The change is associated with petalody of the +stamens and pistils. + +A more frequent change among the monopetalous orders is the duplication +or triplication of the corolla, in consequence of which there appear to +be a series of corollas enclosed one within the other, the lobes of +which generally alternate with one another, but which sometimes are +superposed. This happens occasionally in the primrose (_Primula +acaulis_), and constitutes the variety called by the gardeners "hose in +hose." + +The same condition occurs frequently in some species of _Datura_ and +_Campanula_. + +[Illustration: FIG. 191.--_Campanula rotundifolia_. "Double flowers" +resulting from dialysis and multiplication of the petals.] + +In _Antirrhinum majus_ double flowers of this character sometimes occur; +the outermost corolla is normal, the succeeding ones usually have their +petals separate one from the other; the stamens are sometimes present, +sometimes absent, and at other times petalodic. Similar occurrences may +be met with in labiates and jasmines, and in _Erica hyemalis_. + +Mr. W. B. Hemsley has kindly furnished me with flowers of a similar kind +occurring in wild specimens of _Epacris impressa_,[437] and there are +analogous phenomena in the common honeysuckle (_Lonicera Periclymenum_), +in which three corollas and no stamens often occur. + +This duplication may either be accounted for on the theory of chorisis +above alluded to, or by supposing that the extra corolline whorl is due +to a series of confluent petalodic stamens; that the latter is the true +explanation, in certain cases at least, is shown by some flowers of +_Datura fastuosa_, in which the second corolla was partially staminal in +its appearance, and bore nearly perfect anthers, in addition to the five +ordinary stamens, which were unaltered either in form or position. Some +partially virescent honeysuckle flowers have a similar structure. + +There are other cases of apparent multiplication or duplication, due, +probably, rather to the formation of outgrowths from the petals than to +actual augmentation of their number. These excrescences occur sometimes +on the inner surface of the petals, or of the corolla; at other times on +the outer surface, as in some gloxinias, &c. This matter will be more +fully treated of under the head of hypertrophy and enation. + +=Pleiotaxy of the androecium.=--An increase in the number of whorls in +the stamens is very common, especially in cases where the number of +circles of stamens is naturally large. The augmentation of the number of +stamens is still more frequent where these organs are arranged, not in +verticils, but in one continuous spiral line. + +In _Cruciferae_ there is always an indication of two whorls of stamens, +and this indication is rendered even more apparent in some varieties +accidentally met with. So in _Saponaria_, in _Dianthus_, and other +_Caryophylleae_, three and four verticils of stamens have been met with. +In _Lonicera Periclymenum_ a second whorl of stamens more or less +petalodic sometimes occurs. + +Moquin mentions a variety of _Rubus fruticosus_ in which nearly 900 +petaloid organs existed in the place of the twenty-five or thirty +stamens natural to the plant, the other organs of the flower being in +their ordinary condition, with the exception of the pistil, which did +not attain its full size. Baillon records the occasional existence of +two rows of stamens in _Ditaxis lancifolia_. + +=Increased number of stamens in orchids, &c.=--Various deviations from +the ordinary type of orchid structure have been already alluded to under +the head of displacement, fusion, peloria, substitution, &c., but the +alterations presented by the androecium in this family are so +important in reference to what is considered its natural conformation, +that it seems desirable, in this place, to enter upon the teratological +appearances presented by the androecium in this order, in somewhat +greater detail than usual. The ordinary structure of the flower with its +three sepals, two petals, labellum, column; and inferior ovary, is well +known. Such a conformation would be wholly anomalous and inexplicable +were it not that the real number and arrangement of parts have been +revealed by various workers labouring to the same end in different +fields. Thus, Robert Brown, Link, Bauer, Darwin, and others, paid +special attention to the minute anatomy and mode of distribution of the +vessels; Irmisch, Crueger, Payer, and others, to the evolution of the +flower; Lindley, St. Hilaire, and Reichenbach, to the comparison of the +completed structures in the various genera and species; while the +teratological observers have been numerous, as will be seen from the +selected references cited at the end of this paragraph and in other +places. The result of this manifold study has been a pretty general +agreement that the structure of the order (omitting minor details) is as +follows:--A six-parted perianth in two rows, the outer three (sepals) +generally regular and equal in shape; of the inner three (petals or +tepals) two are regular, and one, the labellum very irregular, +consisting not only of a petal, but of two abortive stamens incorporated +with it. The column is considered to be made up of one perfect and three +abortive stamens, in inseparable connection with three styles. By some, +however, it is supposed that all the stamens are confluent with the +column and none with the lip. + +[Illustration: FIG. 192.--Diagram showing the arrangement of parts in an +orchid flower. According to Crueger, the stamens A 2, A 3, should be +distinct from the lip. The uppermost figure 2 should have been 1. (See +text.)] + +In either case it is admitted that there are six stamens in two rows. +The first row consists of one posterior stamen, which is generally +perfect, and two abortive stamens incorporated with the labellum. The +second row also consists of three stamens, all of which are usually +abortive and inseparable from the column. Traces of them may +occasionally be met with in the form of tubercles or wing-like processes +from the column. In _Cypripedium_, while the ordinary stamen of the +outer row is deficient, two of the inner series are present. The +diagram, fig. 192, will serve to show the arrangement of the parts as +above described. + represents the situation of the stem or axis; on the +opposite side is the bract; between these are placed the sepals, one +posterior or next the axis (incorrectly numbered 2 in the plan), two +lateral 1, 1; next in order follow the petals, 2, 2, 2, two lateral and +somewhat posterior, one larger (the lip), anterior; the outer series of +stamens are represented by A 1, A 2, A 3, the two latter being fused +with the labellum; _a_ 1, _a_ 2, _a_ 3 represent the position of the +inner verticil of stamens, while s, s, s denote the three carpels. It is +foreign to the purpose of this book to detail the varied evidence in +support of this explanation of the homologies of orchid flowers.[438] +All that can be done in these pages is to set forth the evidence +furnished by teratology as to this matter--evidence for the most part +accumulated and recorded without any special reference to any theory of +orchid structure. + +The following details all refer to flowers in which the number of +stamens in orchidaceous plants was increased beyond what is necessary. +They are arranged with reference to the number of adventitious organs, +beginning with those in which the number was smallest, and proceeding +thence to those in which it was greatest. In some cases it has not been +possible to ascertain whether the adventitious organs were really +restorations of the numerical symmetry, substitutions of one part for +another, stamen for petal, &c., or wholly adventitious productions. +Unless otherwise stated, the interpretation put upon the facts thus +recorded is that of the present writer, and not necessarily that of the +original observer. + + Mr. J. T. Moggridge has described and figured a flower of + _Ophrys insectifera_ in which there was a vestige of a second + stamen present, probably one of the inner series fig. 192 + (_a_^2).[439] The same observer also records the presence of a + second anther between the lobes of the normal one. This can + hardly be referred to either of the typical stamens, but would + seem to be a perverted development of the rostellum.[440] + + Roeper is stated by Cramer[441] to have seen a specimen of + _Orchis morio_ with two stamens. + + In a flower of _Habenaria chlorantha_, described by the late + Professor Henslow,[442] the outer three stamens are suppressed, + while two of the inner group are present, as happens normally + in _Cypripedium_. + + A flower of _Cattleya violacea_ afforded a similar + illustration; but in this case only one of the inner stamens + was developed, and this in the form of a small petal, partly + adherent to the column. + + In _Dendrobium normale_, Falconer, not only is the perianth + regular, but the column is triandrous,[443] the three stamens + (according to the diagram of its structure given by Lindley) + pertaining to the outer row. + + In a specimen of _Dendrobium hoemoglossum_ kindly forwarded + from Ceylon by Mr. Thwaites there were three stamens present, + of which one posterior belonged to the outer series A 1, and + two lateral to the inner _a_ 1, _a_ 2, fig. 192. + + M. His observed, several years in succession, some flowers of a + species of _Ophrys_ with three sepals, no lateral petals, one + lip, and three perfect stamens. In this case probably the two + supernumerary stamens were petals which had assumed an + anther-like character. + + Wydler describes a flower of _Ophrys aranifera_ in which one + outer and two inner stamens were present.[444] I have myself + met with three such flowers in the same species. The stamens + present were A 1, _a_ 1, _a_ 2. + + Dr. J. E. Gray exhibited at the Botanical Society of London, in + August, 1843, a specimen of _Ophrys apifera_ with a triandrous + column, the supernumerary anthers belonging, apparently, to the + inner whorl. + + In his 'Catalogue of the Plants of South Kent,' p. 56, tab. iv, + f. 16, the Rev. G. E. Smith describes and figures a flower of + _O. aranifera_ with a triandrous column, seemingly of the same + kind as that spoken of by Dr. Gray. + + Mr. Moggridge met with a triandrous flower in the same species, + and refers the appearance to "a fusion of two flowers, + accompanied by suppression and modification."[445] As, however, + no details are given in support of this opinion, it may be + conjectured that the two additional stamens were members of the + inner whorl _a_ 1, _a_ 2, and thus the conformation would be + the same as in the flowers just mentioned. The figures given by + Mr. Moggridge bear out this latter view, while they lend no + support to the hypothesis advanced by him. Nevertheless, no + decided opinion can be pronounced by those who have not had the + opportunity of examining the flowers in question. + + Alphonse de Candolle[446] figures a flower of _Maxillaria_ in + exactly the same condition, so far as the stamens are + concerned, as in the Ophrys flowers just mentioned. It is + curious to observe that in many of these cases the two lateral + petals are suppressed. + + Von Martius mentions the occurrence of three anthers + (_naturaliter conformatae_) in _Orchis morio_.[447] Richard, + as cited by Moquin-Tandon, Lindley, and others, describes and + figures a peloria of _Orchis latifolia_ with regular triandrous + flowers.[448] + + The writer has examined, in the Royal Gardens at Kew, a flower + of _Cattleya crispa_ in which were three stamens, the central + one normal; the two lateral ones, belonging probably to the + inner whorl, were in appearance like the lateral petals, and + one of them was adherent to the central perfect column. + Duchartre[449] mentions a flower of _Cattleya Forbesii_ in + which there were two labella in addition to the ordinary one, + the column being in its normal condition. From the analogy of + other cases it would appear as if the additional labella in + this instance were the representatives of two stamens of the + outer whorl. Beer likewise has put on record the existence of a + triandrous _Cattleya_.[450] + + A specimen of _Catasetum eburneum_ forwarded by Mr. Wilson + Saunders was normal so far as the sepals and two lateral petals + were concerned, but the anterior petal or labellum was flat and + in form quite like the two lateral ones; the column was normal + and in the situation of the two anterior stamens of the outer + series A 2, A 3, were two labella of the usual form (fig. 156, + p. 291). Perhaps the _Oncidium_ represented at p. 68, fig. 29, + may also be explained on the supposition that the two lateral + lobes of the labellum in this flower were the representatives + of stamens. + + In Fig. 193 is shown the arrangement of parts in a flower of + _Ophrys aranifera_. Here there were three sepals, two lateral + petals, one of which was adherent to the side of the column; + the central labellum was seemingly deficient, but there were + two pseudo-labella placed laterally in the position of the two + antero-lateral stamens of the outer series (A 2, A 3). Within + these was another perfect stamen occupying the position of the + anterior stamen of the inner series (_a_ 3). In another flower + of the same species, gathered at the same time (fig. 194), + there were three sepals not at all different from those of the + normal flower. The three petals next in succession were also, + in form and position, in their ordinary state. In colour, + however, the two upper lateral petals differed from what is + customary, in having the same purplish-brown tint which + characterises the lip. Within these petals, at the upper part + of the flower, there was the ordinary column, and at the + opposite side, alternating with the petals before mentioned, + two additional lip-like petals, one provided with a half-anther + containing a single perfectly formed pollen-mass (A 2, A 3). It + is, perhaps, worthy of notice that the arrangement of the + coloured spots on the true labellum, and that on the + adventitious lips, replacing the two lower of the outer + stamens, were not of a similar character. The supernumerary + lips had the [Greek: pi]-shaped marking which is so common in + this species, while the true lip was, as to its spots, much + more like _O. apifera_. Alternating with this last whorl were + three columns, all apparently perfectly formed and differing + only from the ordinary one in their smaller size and + corresponding to _a_ 1, _a_ 2, _a_ 3. The ovary in this flower + was two-celled, with four parietal placentas, thus giving an + appearance as though there had been a fusion of two or more + flowers associated with suppression and other changes. The + position of the supernumerary organs and the absence of any + positive sign of fusion in the bracts or other part of the + flower, seemed, however, to negative the idea of fusion.[451] + + [Illustration: FIG. 193.--Diagram showing the arrangement of + parts in a malformed flower of _Ophrys aranifera_ (see p. + 384).] + + [Illustration: FIG. 194.--Malformed flower of _Ophrys + aranifera_ with two supernumerary lips and three additional + stamens.] + + A similar illustration, for a knowledge of which the writer is + indebted to the kindness of Professor Asa Gray and Mr. Darwin, + occurred in some specimens of _Pogonia ophioglossoides_ + collected by Dr. J. H. Paine in a bog near Utica, New York. It + will be seen from the following description that these flowers + presented an almost precisely similar condition to those of the + _Ophrys aranifera_ just mentioned. "The peculiarities of these + flowers," writes Professor Gray, "are that they have three + labella, and that the column is resolved into small petaloid + organs. The blossom is normal as to the proper perianth, except + that the labellum is unusually papillose, bearded almost to the + base. The points of interest are, first, that the two accessory + labella are just in the position of the two suppressed stamens + of the outer series, viz. of A2 and A3, as represented in the + diagram, fig. 192; and there is a small petaloid body on the + other side of the flower, answering to the other stamen, A1. + Secondly, in one of the blossoms, and less distinctly in + another, two lateral stamens of the inner series (_a_1 and + _a_2) are represented each by a slender naked filament. There + are remaining petaloid bodies enough to answer for the third + stamen of the inner series and for the stigmas, but their order + is not well to be made out in the dried specimens." It may here + be mentioned that _Isochilus_ is normally triandrous. + + A tetrandrous flower of _Cypripedium_ has also been recorded. + + In _Isochilus_, according to Cruger, there are often five + stamens, and there are several, besides those already + mentioned, in which six more or less perfect stamens have been + seen--of these the following may be taken as illustrations. A + hexandrous flower of _Orchis militaris_ has been recorded by + Kirschleger,[452] and in the accompanying diagram (fig. 195), + from Cramer,[453] of a monstrous flower of _Orchis mascula_, + there is one perfect stamen of the outer row and two lip-like + stamens of the same series, while the inner verticil comprises + one perfect and two abortive stamens. + + [Illustration: FIG. 195.--Diagram of flower of _Orchis mascula_ + with two additional lips, two perfect and two imperfect stamens + (after Cramer).] + + Morren[454] describes some flowers of _Orchis morio_ in which + there were three sepals, three petals, and within the latter + two other ternary series of petals; this would seem to be a + case of petalody of all six stamens. Morren, however, seems to + have considered the additional segments as repetitions of the + corolline whorl, though he describes a central mass as the + column bearing a "_souvenir_ of the anther." Nevertheless, + there is no decisive evidence either in his figure or his + description in support of his opinion as to the nature of the + central mass, which might be a distorted condition of the + styles, or, as is more probable, a rudimentary and irregular + flower. Morren also describes another flower of the same plant + in which there were three sepals, two lateral petals partially + lip-like in aspect, a third labellum normal, two additional + labella representing the two anterior stamens of the outer + whorl, while more or less developed rudiments of the remaining + four stamens also exist. + + While, in most cases, the supernumerary stamens can, by reason + of their relative position, their complete or partial + antheriferous nature, be safely referred to one or other of the + six stamens, making up a typical orchid flower, there are other + specimens in which the additional stamens are altogether + adventitious, and do not admit of reference to the homologue. + Thus it was in a specimen of _Odontoglossum Alexandrae_ examined + by the writer, and in which, within a normally constructed + perianth, there were six columns, all polliniferous, but + arranged in so confused and complicated a manner that it was + impossible to make out any definite relation in their position. + There was nothing to indicate a fusion of flowers, but rather + an extension of the centre of the flower, and consequent + displacement of the stamens, &c. Again, the existence of + adventitious stamens does not necessarily imply the development + of organs usually suppressed, inasmuch as they may result from + the assumption by the lateral petals of staminal + characteristics. + + Nevertheless, as far as teratology is concerned, specimens may + be found in which some or all of the usually suppressed stamens + of _Orchidaceae_ may be found. These stamens may be all perfect + (polliniferous), or, as is more frequently the case, more or + less petal-like. Moreover, when the stamens are petalodic, the + form assumed is usually that of the labellum. + + The presence of stamens in undue numbers in orchids is very + generally, but not always, attended by some coincident + malformation, of which the most frequent is cohesion of two or + more sepals, and consequent displacement or adhesion of one + petal to the side of the column. Petalody of the styles and + median prolification are also sometimes found in association + with an augmented number of stamens. + +[Illustration: FIG. 196.--Increased number of carpels, tulip.] + +[Illustration: FIG. 197.--Fruit of St. Valery apple cut lengthwise.] + +=Pleiotaxy of the gynoecium.=--An increase in the number of whorls of +which the pistil consists is not of very frequent occurrence. Generally +after the formation of the whorl of carpels, the energy of the growing +point ceases, or if by chance it be continued, the result is more +generally the production of a new flower-bud (median prolification) than +the repetition of the carpellary series. It is necessary also to +distinguish between the veritable augmentation of the pistil and the +semblance of it, brought about by the substitution of carpels for some +other organs, as pistillody of the stamens, and even of the segments of +the perianth, is not very unfrequent, as has already been stated under +the head of substitution. Again, the increased number of carpels which +is sometimes met with in such flowers, as _Magnolia_ or _Delphinium_, +where the ovaries are arranged in spiral series, is not strictly +referable to the present category. + +The orange is one of the plants most frequently subject to an +augmentation in the number of carpellary whorls; sometimes this is due +to the stamens assuming the guise of carpels, but at other times the +increase occurs without any alteration in the stamens or other organs. +If the adventitious carpels be exposed, they are covered with yellow +rind, while those portions that are covered by the primary carpels are +destitute of rind. Some varieties of the double tulip are very subject +to a similar change, but, in this case, the petals and the stamens very +frequently become more or less carpellary in their nature. Fig. 196 +represents an increased number of whorls of carpels in the variety +called "rex rubrorum," the segments of the perianth having been removed. + +In the St. Valery apple, already referred to, there is a second whorl of +carpels above the first, a fact which has been made use of to explain +the similar structure of the pomegranate. + +The tomato (_Lycopersicum esculentum_) is another plant in which an +adventitious series is frequently produced, and generally in combination +with the primary series. + +In the Chinese primrose (_Primula sinensis_) a supernumerary whorl is +frequently met with, generally associated with other changes in the +construction and arrangement of the parts of the flower. + +M. de Candolle[455] mentions a flower of _Gentiana purpurea_ with four +carpels in one series, and five others in the circle immediately above +them. Wigand[456] alludes to an instance wherein there was a second pair +of carpels above the first in _Vinca herbacea_. Dr. Sankey has forwarded +flowers of a _Pelargonium_ having a double series of carpels, eight in +the outer row, five in the inner, and this condition is stated to exist +in the flowers of the same plant for two years consecutively. In +_Aquilegia_ I have met with a similar increase in the whorls of +carpels.[457] Meissner records a similar augmentation in _Polygonum +orientale_.[458] + +Wigand[459] describes and figures a flower of _Vinca minor_, in which +there were two carpels intervening between the ordinary pair, and a +similar illustration has been observed by the writer in _Allamanda +cathartica_. Eichler[460] has put on record a similar case in a +capparid. + +Marchand[461] mentions a polycarpellary berberid (_Epimedium +Musschianum_). The supernumerary carpels in this flower were placed on a +short axis, which originated in the axils of the stamens, and as these +latter organs were present in their usual number and position, the +adventitious carpels could not be considered as resulting from a +transformation, or substitution of carpels for stamens. + +Lastly, the instance cited by Dr. Allman[462] in _Saxifraga Geum_ may be +alluded to. Here there was a row of adventitious carpels between the +stamens and pistils, the backs of the carpels being turned towards the +axis of the flowers. Dr. Allman explains the presence of the +supernumerary parts by the supposed production of a whorl of secondary +axes between the stamens and the centre of the flower. These axes are +further supposed to bear imperfect flowers, of which the additional +carpels are the only traces, but this explanation seems forced. + +In addition to the references already cited the following may be given: + + Duchartre, 'Ann. Sc. Nat.,' 4 ser., vii, p. 23 (Tulip). + + Ferrari, 'Hesperides,' pp. 271, 395, 405. Duchartre, 'Ann. Sc. + Nat.,' 4 ser., 1844, vol. i, p. 294. Maout, 'Lecons Element.,' + vol. ii, pp. 488-9. Clos, 'Ann. Sc. Nat.,' 1865, p. 317 + (_Citrus Aurantium_). + + Clos, 'Bull. Soc. Bot. Fr.,' vol. xiii; 'Rev. Bibl.,' p. 75. + Pasquale, 'Reddicont Accad. Sc. Fis. e Math. Napoli.' Octr. + 1866 (_Solanum Lycopersicum_). + +On the general subject of multiplication, in addition to previous +citations, the reader may be referred to A. P. de Candolle, 'Theorie +Element. Bot.,' ed. 3, p. 89. + +=Increased number of flowers in an inflorescence.=--This happens +generally as a result of over luxuriant growth, and scarcely demands +notice here, being rather referable to variation than to malformation. +The increased number of florets in the spikelets of some grasses has +already been alluded to (p. 351). Thus spikelets of wheat occasionally +produce more than the three florets which are proper to them.[463] It +will be remembered that in this as in many other grasses there are +rudimentary florets, and it is no matter for surprise that these florets +should occasionally be fully developed. + +FOOTNOTES: + +[392] 'Bull. Soc. Bot. Fr.,' vol. vii, 1860, p. 587. + +[393] 'Fragment. Phyt. Austral.,' part xx, p. 270. + +[394] 'Bull. Acad. Belg.,' xvi, pt. i, p. 60, "Fuchsia," p. 125, c. ic. + +[395] "Theorie de la feuille," 'Arch. des Sciences Bibl. Univers.,' +1868. + +[396] See Engelmann, 'De Antholysi,' p. 16, section 12. + +[397] Verhandl. des Botanisch. Vereins Brandenburg,' 1859, 1 heft. + +[398] See Henslow. 'Mag. Nat, Hist.' 1832, vol. v, p. 429. + +[399] 'Phytologist,' September, 1857. + +[400] Seemann's 'Journal of Botany,' iv, p. 168, t. 47, f. 3. + +[401] 'Illust. Hortic.,' 1866, misc., p. 97. + +[402] See Fresenius, 'Mus. Senkenb.,' bd. 2, p. 43. Schlechtendal, 'Bot. +Zeit.,' iv, pp. 403, 492, _Veronica tetrandra_. + +[403] 'Flora,' 1865, tab. 6, fig. 8. + +[404] 'Org. Veget.,' t. i, p. 497, pl. 42, f. 3. + +[405] 'El. Ter. Veg.,' p. 354. + +[406] Cited in "Rev. Bibl." of 'Bull. Soc. Bot. Fr.,' 1866, p. 171. + +[407] Loc. cit., 351. + +[408] 'Mem. Acad. Toulous.,' vi, 1862, ex 'Bull. Soc. Bot. Fr.,' "Rev. +Bibl.," vol. ix, 1862. p. 127. + +[409] 'Flora.' 1857. p. 289. + +[410] L. c., p. 354. + +[411] Giraud, 'Ed. Phil. Mag.,' Dec., 1839. + +[412] See _Cerasus Caproniana_, D. C. 'Plant. Rar. Hort. Genev.,' tab. +18. + +[413] Nees, 'Linnaea,' v, p. 679, tab. 11 (_Schoenodorus_). + +[414] 'Gard. Chron.,' 1852, p. 452. + +[415] See Cramer, 'Bildungsabweich.' pp. 16, 24. + +[416] 'Linnaea,' 1842, p. 389, c. ic. + +[417] Seemann's 'Journal of Botany,' 1867, vol. v, p. 158. + +[418] Cramer, 'Bildungsabweich,' p. 66, _Astrantia major_, _Eryngium_, +to which may be added _Daucus_, _Heracleum_, &c. + +[419] See also Reinwardt, 'Nov. Act. Acad. Nat. Cur.,' 12, 1, 37; and +Masters, 'Journ. Linn. Soc.,' vi, p. 24. + +[420] 'Organog. Veget.,' tab. 53. + +[421] 'Missbild.,' p. 206. + +[422] Ehrenberg, 'Flora,' 1846, p. 704. + +[423] 'Flora,' 1860, tab. 7. + +[424] 'Ann. Sc. Nat.,' 3 ser., t. x, p. 207. + +[425] 'Mem. Mus.,' xii. t. 17. + +[426] 'Nov. Act. Acad. Nat. Cur.,' xv, tab. xxviii, f. 3; 'Bot. Mag.,' +t. 1622. "Caryophyllus spicam frumenti referens." A similar malformation +in _Dianthus barbatus_ is not uncommon. It has lately been introduced +into gardens under the name of _Dianthus_ "_mousseux_," but is not +likely to find favour with gardeners. + +[427] 'Bull. Soc. Bot. France,' t. vi, 1859. p. 268. + +[428] Weber, 'Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,' 1860. + +[429] 'Bot. Zeit.,' 1848, p. 217. + +[430] 'De Anthol.,' p. 17, Sec. 12. + +[431] 'Linnaea,' vol. ii, 1827, p. 85. + +[432] 'De Antholysi,' p. 17, tab. iii, f. 15, 16; Weinmann, 'Phytanth. +iconogr.,' nro. 292. + +[433] See Hildebrand, 'Bot. Zeit.,' 1862, p. 209, tab. viii; Cramer, +'Bildungsabweich.,' p. 7, tab. xiii; Engelmann, 'De Antholysi,' p. 18, +&c. For similar changes in _Gagea arvensis_ see Wirtgen, 'Flora,' 1838, +t. xxi. p. 350, and 'Flora.' 1846, p. 353. Some of these are cases of +synanthy. + +[434] Schlechtendal, 'Bot. Zeit.,' xx, 1862, p. 301. + +[435] 'Phil. Bot.,' Sec. 126. + +[436] C. Morren, 'Bull. Acad. Belg.,' xix, part ii, p. 17. + +[437] 'Seemann's Journal of Botany,' iii, p. 354. + +[438] On this point the reader will find an excellent summary in +Lindley's 'Vegetable Kingdom,' cd. iii, p. 183_a_, and in Darwin, +'Fertilisation of Orchids,' p. 292. See also Crueger,'Journ. Linn. Soc.,' +t. viii, p. 134. + +[439] 'Seemann's Journal of Botany,' vol. iv, p. 168, tab. 47. + +[440] Ibid., t. iv. 1866, p. 168, t. xlvii, f. 1. + +[441] 'Bildungsabweich,' p. 8; see also 'Bot. Zeit.,' 1852, p. 425. + +[442] 'Journ. Linn. Soc.,' t. ii, p. 104. tab. 1, fig. B. + +[443] Lindl., "Orchid. Ind.," 'Jour. Linn. Soc.,' iii, p. 9. + +[444] 'Arch. Bot.,' ii, p. 300, tab. xvi, f. 11. + +[445] 'Seemann's Journal of Botany,' v, p. 318, tab. lxxii, figs. A 4, 4 +_a_. + +[446] "Monstr. Veg.," in 'Neue Denkschrift,' p. 17, tab. vii. + +[447] 'Flora,' t. viii, 1825, p. 736. + +[448] 'Mem. Soc. d'Hist. Nat.,' ii, 1, p. 212, tab. iii. + +[449] 'Bull. Soc. Bot. Fr.,' t. vii, 1860, p. 26. + +[450] 'Beitr. Morphol. und Biol. Orchid.,' quoted by Cramer; +'Bildungsabweich,' p. 9. + +[451] Masters, 'Journ. Linn. Soc.,' viii, p. 207. See also Rodigas, +'Bull. Soc. Bot. Belg.,' iv, p. 266, for similar changes in _Cypripedium +Hookerae_. + +[452] Kirschleger, 'Flora,' 1844, p. 131. + +[453] 'Bildungsabweich,' p. 11, tab. xiv, f. 3. + +[454] 'Bull. Acad. Roy. Belg.,' t. xix, part 2, p. 171. + +[455] 'Organogr. Veget.,' t. i, p. 509, tab. 40, figs. 6, 7. + +[456] 'Flora,' 1856, p. 715. + +[457] 'Linn. Trans.,' t. xxiii, p. 364, tab. 34, fig. 5. + +[458] 'Monog. Polygon,' pl. 3, K. f. 12. + +[459] 'Flora,' 1856, tab. viii. + +[460] Ibid., 1865, tab. ix, f. 6. + +[461] 'Adansonia,' vol. iv, 1864, p. 127. + +[462] 'Ann. Nat. Hist.,' 1845, vol. xvi, p. 126. + +[463] See Schlechtendal, 'Bot. Zeit.,' t. xviii, p. 381 (_Triticum_); +also 'Flora,' t. xiv, 1831, p. 5 (_Avena_). + + + + +PART II. + +DIMINISHED NUMBER OF ORGANS. + + +A diminution in the number of parts is generally due to suppression, +using that word as the equivalent of non-development. It corresponds +thus in meaning with the _Fehlschlagen_ of the Germans, the _avortement +complete_ of Moquin and other French writers. It differs from atrophy, +or partial abortion, inasmuch as the latter terms apply to instances +wherein there has been a partial development, and in which evolution has +gone on to a certain extent, but has, from some cause or other, been +checked. These cases will be found under the head of diminished size of +organs. As the word abortion is used by different authors in different +ways, it is the more necessary to be as precise as possible in the +application of the term. In the present work abortion is used to apply +to cases wherein parts have been formed, but wherein growth has been +arrested at a certain stage, and which, therefore, have either remained +_in statu quo_, while the surrounding parts have increased, or have, +from pressure or other causes, actually diminished in size. + +In practice, however, it is not always possible to discriminate between +those instances in which there has been a true suppression, an absolute +non-development of any particular organ, and those in which it has been +formed, and has grown for a time, but has afterwards ceased to do so, +and has been gradually obliterated by the pressure exercised by the +constantly increasing bulk of adjacent parts, or possibly has become +incorporated with them. In the adult flower the appearances are the +same, though the causes may have been different. + + + + +CHAPTER I. + +SUPPRESSION OF AXILE ORGANS. + + +Absolute suppression of the main axis is tantamount to the non-existence +of the plant, so that the terms "acaulescent," "acaulosia," etc, must be +considered relatively only, and must be taken to signify an atrophied or +diminished size of the stem, arising from the non-development of the +internodes. + +The absence of lateral branches or divisions of the axis is of frequent +occurrence, and is dependent on such causes as the following:--deficient +supply of nutriment, position against a wall or other obstacle, close +crowding of individual plants, too great or too little light, too rich +or too poor a soil, &c. + +Probably the absence of the swollen portion below the flower in the case +of many proliferous roses, double-flowered apples, as already referred +to, may be dependent on the non-development of the extremity of the +peduncle or flower-stalk. Thus, in a double-flowered apple recently +examined, there was a sort of involucel of five perfect leaves, then +five sepals surrounding an equal number of petals, numerous stamens, and +five styles, but not a trace of an expanded axis, nor of any portion of +the carpels, except the styles. The views taken as to the nature of this +and similar malformations must depend on the opinion held as to the +nature of inferior pistils, and on the share, if any, that the expanded +axis takes in their production. As elsewhere said, the evidence +furnished by teratology is conflicting, but there seems little or +nothing to invalidate the notion that the end of the flower-stalk and +the base of the calyx may, to a varying extent, in different cases, +jointly be concerned in the formation of the so-called calyx-tube and of +the inferior ovary. Obviously it is not proper to apply to all cases +where there is an inferior ovary the same explanation as to how it is +brought about. + +As these pages are passing through the press, M. Casimir de Candolle has +published a different explanation as to the nature of the hip of the +rose, having been led to his opinion by the conclusion that he has +arrived at, that the leaf is to be considered in the light of a +flattened branch, whose upper or posterior surface is more or less +completely atrophied. + +According to M. de Candolle, the calyx-tube, in the case of the rose, is +neither a whorl of leaves, nor a concave axis in the ordinary sense in +which those terms are used, but is rather to be considered as a +ring-like projection from an axis arrested in its ulterior development. +The secondary projections from the original one correspond to an equal +number of vascular bundles, and develope into the sepals, petals, +stamens, and ovaries. If these organs remained in a rudimentary +condition, the tube of the calyx would be reduced to the condition of a +sheathing leaf. The rose flower, then, according to M. de Candolle, may +be considered as a sheathing leaf, whose fibro-vascular system is +complete, and from which all possible primary projections are +developed.[464] + +If, as M. de Candolle considers, the leaf and the branch differ merely +in the fact that the vascular system is complete in the latter, and +partly atrophied in the former, it would surely be better to consider +the "calyx-tube" of the rose as a concave axis rather than as a leaf, +seeing that he admits the fibro-vascular system to be complete in the +case of the rose. + +With reference to this point the reader is referred to Mr. Bentham's +account of the morphology and homologies of the _Myrtaceae_ in the +'Journal of the Linnean Society,' vol. x, p. 105. See also _ante_, pp. +71, 77. + +Some doubts also exist as to the nature of the beak or columella of such +fruits as those of _Geraniaceae_, _Malvaceae_, _Umbelliferae_, +_Euphorbiaceae_, &c. The nature of the organ in question may probably be +different in the several orders named; at any rate the subject cannot be +discussed in this place, and it is mentioned here because, now and then, +it happens that the organ in question is completely wanting, and hence +affords an illustration of suppression. + +FOOTNOTES: + +[464] 'Theorie de la feuille.' p. 24. + + + + +CHAPTER II. + +SUPPRESSION OF FOLIAR ORGANS. + + +This subject may be considered, according as the separate leaves of the +stem or of the flower are affected, and according as either the number +of members of distinct whorls, or that of the whorls themselves, is +diminished. + +The terms aphylly, meiophylly, and meiotaxy may be employed, according +as the individual leaves are altogether wanting, or with reference to +the diminished number of parts in a whorl, or a decrease in the +verticils. + +=Aphylly.=--Entire suppression of the leaves is a rare phenomenon. Under +ordinary circumstances it occurs in most _Cactaceae_, in some of the +succulent Euphorbias, and other similar plants, where the epidermal +layers of the stem fulfil the functions of leaves. But even in these +plants leaf-like organs are present in some stage or another of the +plant's life. + +Partial suppression of the leaf occurs sometimes in compound leaves, +some or other of the leaflets of which are occasionally suppressed. +Sometimes, as Moquin remarks, it is the terminal leaflet which is +wanting, when the appearance is that of _Cliffortia_, at other times the +lateral leaflets are deficient, as in _Citrus_ or _Phyllarthron_. +_Ononis monophylla_ and _Fragaria monophylla_ may be cited as instances +of the suppression of the lateral leaflets. If the blade of the leaf +disappears entirely, we have then an analogous condition to that of the +phyllodineous acacias. + +With reference to the strawberry just mentioned, Duchesne, 'Hist. Nat. +Frais.,' p. 133, says that this was a seedling raised from the _fraisier +des bois_, and the characters of which were reproduced by seed, and have +now become fixed. The monophyllous condition has been considered to be +the result of fusion of two or more leaflets, but however true this may +be in some cases, it is not the case with this strawberry. M. Paillot +states that he has found the variety in a wild state.[465] + +In like manner varieties of the following plants occur with simple +leaves, _Rosa berberifolia_ (_Lowea_), _Rubus Idaeus_, _Robinia +pseudacacia_, _Fraxinus excelsior_, _Sambucus nigra_, _Juglans nigra_, +&c. + +In one instance seen by the writer every portion of the leaf of a rose +was deficient, except the stipules and a small portion of the petiole. +(See abortion.) + +=Meiophylly.=--A diminished number of leaves in a whorl occasionally +takes place; thus, in some of the _Stellatae_, and frequently in _Paris +quadrifolia_, the number of leaves in the verticil is reduced. Care must +be exercised in such instances that an apparent diminution arising from +a fusion of two or more leaves be not confounded with suppression. + +=Meiophylly of the calyx or perianth.=--A lessened number of sepals is +not a very common occurrence among dicotyledonous plants. Seringe +figures a proliferous flower of _Arabis alpina_ with two sepals only, +and a similar occurrence has been noticed in _Diplotaxis tenuifolia_. + +In _Cattleya violacea_ the writer has met with a flower in which the +uppermost sepal was entirely wanting, while two of the lateral petals +were fused together. Moquin records that in some of the flowers of +_Chenopodiaceae_, in which the inflorescence is dense, a suppression of +two or three sepals sometimes occurs. The species mentioned are _Ambrina +ambrosioides_, _Chenopodium glaucum_, and _Blitum polymorphum_. + +=Meiophylly of the corolla.=--Suppression of one or more petals is of +more frequent occurrence than the corresponding deficiency in the case +of the sepals. Among _Caryophyllaceae_ imperfection as regards the +numerical symmetry of the flower is not uncommon, as in species of +_Cerastium_, _Sagina_, _Dianthus_, &c. In _Ranunculaceae_ the petals are +likewise not unfrequently partially or wholly suppressed. A familiar +illustration of this is afforded by _Ranunculus auricomus_, in which it +is the exception to find the corolla perfect.[466] Some varieties of +_Corchorus acutangulus_ in west tropical Africa are likewise subject to +the same peculiarity. Amongst _Papilionaceae_ absence of the carina or of +the alae is not uncommon, as in _Trifolium repens_, _Faba vulgaris_, &c. + +Moquin relates a case of the kind in the haricot bean, in which the +carina was entirely absent, and another in the pea, where both carina +and alae were missing, thus reducing the flower to the condition that is +normal in _Amorpha_ and _Afzelia_. Suppression of the upper lip in such +flowers as _Calceolaria_ has been termed by Morren "apilary." + +In _Orchidaceae_ entire absence of the labellum, frequently without any +other perceptible change, is of common occurrence. The writer has seen +numerous specimens of the kind in _Ophrys apifera_ and _O. aranifera_; +also in _Dendrobium nobile_, _AErides odoratum_, _Cypripedium villosum_, +_Listera ovata_, &c. Morren[467] mentions analogous deficiencies in +_Zygopetalum maxillare_, _Calanthe_ sp., and _Cattleya Forbesii_. In +most of these there was also a fusion of the two lower sepals, which +were so twisted out of place as to occupy the situation usually held by +the labellum. At the same time the column was partially atrophied. To +this deficiency of the lip the author just quoted proposed to apply the +term acheilary, [Greek: a-cheilarion]. Mr. Moggridge has communicated to +the author an account of certain flowers of _Ophrys aranifera_, in which +the petals were deficient, sometimes completely, at other times one or +two only were present. + +=Meiophylly of the androecium.=--Suppression of one or more stamens, +independently of like defects in other whorls, is not uncommon, even as +a normal occurrence, _e.g._ in _Carlemannia_, where the flower, though +regular, has only two stamens, and other similar deficiencies are common +in Dilleniads. + +Seringe relates the occurrence of suppression of some of the stamens in +_Diplotaxis tenuifolia_,[468] St. Hilaire in _Cardamine hirsuta_, others +in _C. sylvatica_. + +In _Caryophyllaceae_ suppression of one or more stamens has been observed +in _Mollugo cerviana_, _Arenaria tetraquetra_, _Cerastium_, &c.[469] +Among violets the writer has observed numerous flowers in which two or +three stamens were suppressed. Chatin[470] alludes to a similar +reduction in _Tropaeolum_, while in flowers that are usually didynamous +absence of two or more of the stamens is not unfrequent, _e.g._ in +_Antirrhinum_, _Digitalis_, while in a flower of _Catalpa_ a solitary +perfect stamen, and a complete absence of the sterile ones usually +present, have been observed. This might have been anticipated from the +frequent deficiencies in the staminal whorl in these plants under what +are considered to be normal conditions. Reduction of the staminal whorl +is also not unfrequent in _Trifolium repens_ and _T. hybridum_, and has +been seen in _Delphinium_, &c.[471] + +=Meiophylly of the gynoecium.=--Numerical inequality in the case of +the pistil, as compared with the other whorls of the flower, is of such +common occurrence, under ordinary circumstances, that in some text-books +it is looked on as the normal condition, and a flower which is isomerous +in the outer whorls is by some writers not considered numerically +irregular if the number of the carpels does not coincide with that of +the other organs. + +But in this place it is only necessary to allude to deviations from the +number of carpels that are ordinarily found in the particular species +under observation. As illustrations the following may be +cited:--_Arenaria tetraqueta_, which has normally three styles, and a +six-valved capsule, has been seen with two styles, and a four or +five-valved capsule. Moquin relates an instance in _Polygala vulgaris_ +where there was but a single carpel, a condition analogous to that which +occurs normally in the allied genus _Mozinna_. _Reseda luteola_ +occasionally occurs with two carpels only, while Aconites, Delphiniums, +Nigellas, and Paeonies frequently experience a like diminution in their +pistil. + +In a flower of _Papaver Rhaeas_ the writer has recently met with an ovary +with four stigmas and four parietal placentae only, and to Mr. +Worthington Smith he is indebted for sketches of crocus blooms with two, +and in one instance only a solitary carpel. + +Moquin cites the fruit of a wild bramble (_Rubus_) in which all the +little drupes which go to make up the ordinary fruit were absent, except +one, which thus resembled a small cherry. In _Crataegus_ the pistil is +similarly reduced to a single carpel, as in _C. monogyna_. + +The writer has on more than one occasion met with walnuts (_Juglans_) +with a single valve and a single suture.[472] If the ovary of _Juglans_ +normally consisted of two valvate carpels, the instances just alluded to +might possibly be explained by the suppression of one carpel, but the +ovary in _Juglans_ is at first one-celled according to M. Casimir de +Candolle. + +Among monocotyledons _Convallaria majalis_ may be mentioned as very +liable to suffer diminution in the number of its carpels, either +separately or in association with other changes.[473] + +=Meiophylly of the flower as a whole.=--In the preceding sections a +reduction in the parts of each individual whorl has been considered +without reference to similar diminution in neighbouring verticils. It +more commonly happens, nevertheless, that a defect in one series is +attended by a corresponding imperfection in adjoining ones. Thus +trimerous fuchsias and tetramerous jasmines may frequently be met with, +and Turpin describes a tetramerous flower of _Cobaea scandens_. Perhaps +monocotyledonous plants are more subject to this numerical reduction of +the parts of several verticils than are other flowering plants. Thus, in +both _Lilium lancifolium_ and _L. auratum_ the writer has frequently met +with pentamerous flowers. In _Convallaria maialis_ a like deviation not +unfrequently occurs.[474] M. Delavaud has recorded a similar occurrence +in a tulip.[475] + +Dimerous crocuses may also sometimes be met with. In one flower of this +nature the segments of the perianth were arranged in decussating pairs, +and the four stamens were united by their filaments so as to form two +pairs. + +M. Fournier mentions something of the same kind in the flower of an +_Iris_.[476] + +Orchids seem peculiarly liable to the decrease in the number of their +floral organs. Prillieux[477] mentions a flower of _Cattleya +amethystina_ wherein each whorl of the perianth consisted of two +opposite segments. + +The same observer has put on record instances of a similar kind in +_Epidendrum Stamfordianum_. In one flower of the last-named species the +perianth consisted of one sepal only, and one lip-like petal placed +opposite to it.[478] Morren[479] describes a flower of _Cypripedium +insigne_, in which there were two sepals and two petals. Of a similar +character was the flower found by Mr. J. A. Paine, and described in the +following terms by Professor Asa Gray in the 'American Journal of +Science,' July, 1866:--"The plant" (_Cypripedium candidum_) "bears two +flowers: the axillary one is normal; the terminal one exhibits the +following peculiarities. The lower part of the bract forms a sheath +which encloses the ovary. The labellum is wanting; and there are two +sterile stamens, the supernumerary one being opposite the other, _i.e._ +on the side of the style where the labellum belongs. Accordingly the +first impression would be that the labellum is here transformed into a +sterile stamen. The latter, however, agrees with the normal sterile +stamen in its insertion as well as in shape, being equally adnate to the +base of the style. Moreover, the anteposed sepal is exactly like the +other, has a good midrib and an entire point. As the two sterile stamens +are anteposed to the two sepals, so are the two fertile stamens to the +two petals, and the latter are adnate to the style a little higher than +the former. The style is longer than usual, is straight and erect; the +broad, disciform stigma therefore faces upwards; it is oval and +symmetrical, and a light groove across its middle shows it to be +dimerous. The placentae, accordingly, are only two. The groove on the +stigma and the placentae are in line with the fertile stamens. + +Here, therefore, is a symmetrical and complete, regular, but dimerous +orchideous flower, the first verticil of stamens not antheriferous, the +second antheriferous, the carpels alternate with these; and here we have +clear (and perhaps the first direct) demonstration that the orchideous +type of flower has two stamineal verticils, as Brown always insisted." + +[Illustration: FIG. 198.--Regular dimerous flower of _Calanthe +vestita_.] + +[Illustration: FIG. 199.--Regular dimerous flower of _Odontoglossum +Alexandrae_.] + +Dr. Moore, of Glasnevin, kindly forwarded to the writer a flower of +_Calanthe vestita_ (fig. 198), in which there were two sepals only, +anterior and posterior, and two petals at right angles to the two +sepals. The lip was entirely wanting, but the column and ovary were in +their usual condition. In _Odontoglossum Alexandrae_ a similar reduction +of parts has been observed by the author (fig. 199). + +It is curious to observe in these flowers how precisely one sepal +occupies the position of the labellum, and how the lateral petals are +displaced from the position they usually occupy, so as to form a regular +flower, the segments of which decussate, thus giving rise to a species +of regular peloria. + +The genus _Maelenia_ was established on a malformed flower of _Orchis_ of +similar character to those above mentioned. + +=Meiotaxy of the calyx.=--As already mentioned, this term is here +employed to denote those illustrations in which entire whorls are +suppressed. Complete deficiency of the calyx in a dichlamydeous flower +seems seldom or ever to occur; the nearest approach to it would be in +those cases where the calyx is, as it is termed, "obsolete," but here it +is chiefly the limb of the calyx which is atrophied, the lower portion +being more or less adherent to the ovary. In what are termed +monochlamydeous flowers both calyx and corolla are wanting, as in +_Salicineae_ and many other orders. + +=Meiotaxy of the corolla.=--Deficiency of the entire corolla occurs in +conjunction with similar reductions in other organs, or as an isolated +phenomenon in the many apetalous varieties of plants recorded in books. +Deficiency of the corolla was observed in _Campanula perfoliata_ and +_Ruellia clandestina_ by Linne, who calls such blooms _flores +mutilati_.[480] Drs. Hooker and Thomson relate a similar occurrence in +_Campanula canescens_ and _C. colorata_. Some plants seem as a normal +occurrence to produce flowers of different construction, and are hence +termed dimorphic, as in many _Malpighiaceae_, _Violaceae_, _Oxalidaceae_, +in some of the flowers of which the petals are altogether wanting, while +in others the corolla is developed as usual. This deficiency of the +corolla is frequently, but not invariably, associated with an increased +fertility. Thus, in some violets the flowers produced in summer, and in +which the petals are either entirely suppressed or are more or less +atrophied, are always fertile, while the blossoms developed in spring, +and in which the petals are always present, are much less fertile. In +_Oxalis Acetosella_ there are two forms of flower, the one with, the +other without, petals, but both seem equally fertile. Linne remarks that +many plants which, in warm latitudes, produce a corolla, do not do so +when grown in colder climates. Thus, certain species of _Helianthemum_ +are apetalous in Lapland. In the Pyrenees, according to Bentham, the +flowers of _Ajuga iva_ are constantly deprived of their corolla.[481] + +Apetalous flowers have been noted most frequently in the following +plants: + + Aconitum, sp. pl.! + Cardamine impatiens. + Cheiranthus Cheiri! + Viola odorata! + Cerastium vulgatum! + Alsine media. + Stellaria. + Lychnis dioica! + Dianthus barbatus, and other Caryophylleae. + Helianthemum, sp.! + Oxalis Acetosella. + Balsamineae. + Malpighiaceae. + Rosa centifolia. + arvensis! + Crataegus! + Medicago lupulina. + Melilotus officinalis. + Ononis minutissima. + Saxifraga longifolia. + Verbascum Thapsus. + Ajuga iva. + Teucrium Botrys. + Lamium purpureum! + amplexicaule. + Polemonium caeruleum. + Campanula, sp. pl.! + Ruellia clandestina. + Lonicera Periclymenum! + Tradescantia, sp.! + Hymenocallis. + + The following references apply some to apetalous and others to + dimorphic flowers, but it must be remembered that the latter + plants are not necessarily wanting in petals or stamens, &c., + though the functional activity of the parts may be impaired: + + A. de Jussien, 'Monogr. Malpigh.,' pp. 82, 334. Torrey, 'Fl. + New York,' i, p. 428. Hooker and Thomson, 'Journ. Linn. Soc.,' + ii, p. 7, Guillemin, 'Archiv. de Botan.,' i, p. 412. Michalet, + 'Bull. Soc. Bot. Fr.,' vii. p. 465. Mueller, 'Bot. Zeit.,' 1857, + p. 729. 'Natural History Review,' July, 1862, p. 235. + +=Meiotaxy of the androecium.=--Complete suppression of the stamens +occurs normally in the female flowers of unisexual plants, and, as an +accidental occurrence, is not very uncommon. _Erica Tetralix_ is one of +the plants in which this is said to happen. The variety _anandra_ is +said to have been known in France since 1635. Cornuti speaks of it in +his 'Enchiridion.' In 1860 M. du Parquet discovered it in peaty woods +near Nangis (Seine et Marne). + +Many _Umbelliferae_, such as _Trinia vulgaris_, present a like +deficiency, while it is of common occurrence among _Rosaceae_ and +_Pomaceae_. In the latter group the St. Valery apple, so often referred +to, is an illustration. To obtain fruits from this variety it is +necessary to apply pollen from another flower, a proceeding made the +occasion of festivity and rejoicing by the villagers in some parts of +France. In some of the _Artemisias_, especially in _Artemisia +Tournefortiana_, all the florets have been noticed to be female, owing +to the suppression of the stamens, and this suppression is associated +with a change in the form of florets.[482] Mr. Moggridge has +communicated to the author flowers of _Thymus Serpyllum_ from a plant in +which all the stamens were deficient, the flower being otherwise normal. + +M. Dupont has given a list of nineteen species of _Chenopodiaceae_ in +which female flowers are occasionally produced, owing to the entire +suppression of the staminal whorl.[483] + +Flowers the subjects either of regular or irregular peloria, _q. v._, +are often destitute of some or all their stamens, _e.g._ _Calceolaria_, +_Linaria_, _Viola_, &c., while in cases of synanthy suppression of some +of the parts of the flower, and specially of the stamens, is of very +common occurrence. + +Suppression of the androecium as a teratological occurrence has been +most frequently noticed in the following plants, omitting members of +those families whose floral construction is normally incomplete in the +majority of instances, and exclusive also of cases of substitution. See +also under Heterogamy. + + Ranunculus Ficaria! + auricomus! + bulbosus! + Cruciferae, sp. pl. + Violaceae, sp. pl. + Honckenya peploides. + Stellaria. + Caryophyllaceae, sp. pl. + Malpighiaceae, sp. pl. + Tropaeolum majus! + Fragaria vesca! + Rubus, sp. + Pyrus Malus. + Agrimonia vulgaris. + Rosaceae, sp. pl. + Trifolium hybridum. + repens. + Umbelliferae, sp. pl. + Onagraceae, sp. pl. + Hippuris vulgaris. + Callitriche vernalis. + autumnalis. + Lonicera Periclymenum. + Erica Tetralix. + Thymus Serpyllum. + Calceolaria. + Compositae, sp. pl. + Chenopodiaceae, sp. pl. + Stratiotes aloides. + +=Meiotaxy of the gynoecium.=--Complete suppression of the pistil is of +more frequent occurrence than that of the stamens, hence more flowers +become accidentally unisexual by suppression of the pistil than by +deficiency of the stamens. + +In many _Umbelliferae_, e.g. _Torilis Anthriscus_, _Cicuta virosa_, the +central flowers are often male, owing to the suppression of the pistil. +In many double flowers, owing to the excessive multiplication of +petaloid stamens, the pistil is suppressed, in which cases it often +happens that the flower is depressed in the centre, as in some garden +varieties of _Ranunculus_. Schlechtendal, in describing a flower of +_Colchicum autumnale_, in which the perianth was virescent, says that, +although the stamens were present, the pistil was absent. + +In proliferous flowers the pistil is often completely defective, its +place being occupied by the adventitious bud or axis. + +As in other cases of like nature, suppression of the pistil is very +frequently consequent on fusion of flowers or other changes. Thus Morren +relates an instance of synanthy in the flowers of _Torenia scabra_, +accompanied by resorption or disappearance of some parts and spiral +torsion of others. The pistil was entirely absent in this instance.[484] + +M. Gaetano Licopoli places on record an instance where the petals and +carpels of _Melianthus major_ were suppressed.[485] + +On the whole, the pistil seems less subject to changes of this character +than the androecium. + +Suppression of the pistil has been most frequently recorded in flowers +(normally bisexual) of-- + + Ranunculus! + Aconitium! + Delphinium! + Paeonia. + Caryophylleae! + Umbelliferae. + Trifolium repens. + hybridum. + Compositae, sp. pl. + Datura. + Torenia asiatica. + Colchicum autumnale. + +=Suppression of ovules,--abortion of seeds.=--The two cases are taken +together, as the effects are similar, though it must be remembered that +in the one case the ovules at any rate have been formed, but their +development has been arrested, while in the other they have never +existed. The precise cause that has determined the absence of seed +cannot in all cases be ascertained in the adult condition, hence it is +convenient to treat the two phenomena under one head. + +Many plants in other than their native climates either produce no fruit +at all, or the fruits that are produced are destitute of seed, _e.g._ +_Musa_, _Artocarpus_, &c. Some of the cultivated varieties of the grape +and of the berberry produce no seeds. + +Suppression or abortion of the seed is frequently associated with the +excessive development either in size or number of other portions of the +plant, or with an altered condition, as when carpels become foliaceous +and their margins detached. Hybridisation and cross fertilisation are +also well-known agents in diminishing the number and size of seeds. + +=Meiotaxy of the parts of the flower in general.=--In the preceding +sections suppression has been considered as it affected individual +members of a whorl or separate whorls. It rarely happens, however, that +the suppression is limited in this way. More generally several of the +parts of the flower are simultaneously affected in the same manner. + +A few illustrations are all that is necessary to give as to this point. + +One of the most familiar instances is that of the cauliflower or +broccoli, where the common flower-stalk is inordinately thickened and +fleshy, while the corolla and inner parts of the flower are usually +entirely suppressed; the four sepals can, however, generally be +detected. + +Maximowicz describes a _Stellaria_ (_Kraschenikovia_) in which the upper +flowers are male only, while the lower ones, which ultimately become +buried in the soil, have neither petals, stamens, nor styles, but the +walls of the capsule are fleshy, and enclose numerous seeds.[486] + +Kirschleger[487] mentions a variety of _Lonicera Caprifolium_, which was +not only destitute of petals but of stamens also. + +In some species of _Muscari_ and _Bellevalia_ the uppermost flowers of +the raceme show more or less complete suppression of almost all the part +of which the flower normally consists. In those cases where an imperfect +perianth exists, but in which the stamens and pistils are entirely +suppressed, Morren applies the term Cenanthy, [Greek: kenos], empty. + +=Complete suppression of the flower.=--It is not necessary in this place +to allude to that deficient production of flowers characteristic of what +is termed by gardeners a "sky bloomer." In such plants often the +requisite conditions are not complied with, and the skill of the +gardener is shown in his attempt to discover and allow the plant to +avail itself of the necessary requirements. We need here only allude to +those instances in which provision is made for the production of +flowers, and yet they are not produced. A good illustration of this is +afforded by the feather-hyacinth, _Hyacinthus comosus_, in which the +flowers are almost entirely suppressed, while the pedicels are +inordinately increased in number, and their colour heightened. Something +similar occurs in several allied species, and in _Bowiea volubilis_. The +wig plant (_Rhus Cotinus_) affords another illustration of the same +thing. Some tendrils also owe their appearance to the absence of +flowers, being modified peduncles; proofs of this may frequently be met +with in the case of the vine. + +In _Lamium album_ I have seen one of the verticillasters on one side of +the stem completely wanting, the adjacent leaf being, however, as fully +formed as usual. + +=General remarks on suppression.=--On comparing together the various +whorls of the flower in reference to suppression, and, it may be added, +to atrophy, we find that these phenomena occur most rarely in the calyx, +more frequently in the corolla, and very often in the sexual +organs and seeds; hence it would seem as if the uppermost and most +central organs, those most subject to pressure and latest in date of +development--formed, that is, when the formative energies of the plant +are most liable to be exhausted--are the most prone to be suppressed or +arrested in their development. When the plants in which these +occurrences happen most frequently are compared together, it may be seen +that partial or entire suppression of the floral envelopes, calyx, and +corolla, is far more commonly met with in the polypetalous and +hypogynous groups than in the gamopetalous or epigynous series. + +The orders in which suppression (speaking generally) occurs most often +as a teratological occurrence are the following:--_Ranunculaceae_, +_Cruciferae_, _Caryophyllaceae_, _Violaceae_, _Leguminosae_, _Onagraceae_, +_Jasminaceae_, _Orchidaceae_. It will be observed that these are all +orders wherein suppression of the whole or part of the outer floral +whorls takes place in certain genera as a constant occurrence. + +Again, it may be remarked that many of these orders show a tendency +towards a regular diminution of the assumed normal number of their +parts; thus, among _Onagraceae_, _Circeia_ and _Lopezia_ may be referred +to, the former normally dimerous, the latter having only one perfect +petal. So in fuchsias, a very common deviation consists in a trimerous +and rarely a dimerous symmetry of the flower. + +Although, if the absolute number of genera or orders be counted, there +appears to be little difference in the frequency of the occurrence of +suppression in irregular flowers as contrasted with regular flowers, yet +if the individual instances could be counted in the two groups +respectively it would be found that suppression is more common among +irregular than in regular flowers. Thus, the number of individual +instances of flowers in which the perianth is defective is comparatively +large among _Violaceae_, _Leguminosae_, and _Orchidaceae_. This statement +hardly admits of precise statistical proof; still, it is believed that +any observer who pays attention to the subject must come to the same +conclusion. This is but another illustration of the fact that conditions +which are abnormal in one plant constitute the natural arrangement in +others. + +As to the suppressions that occur in the case of the sexual organs, and +the relations they bear to dimorphism, diclinism, &c., but little stress +has been laid on them in this place, because their chief interest is in +a physiological point of view, and is treated of in the writings of +Mohl, Sprengel, Darwin, Hildebrand, and others. All that need be said +here is, that teratology affords very numerous illustrations of those +intermediate conditions which are also found, under natural +circumstances, between the absolutely unisexual flowers, male or female, +and the structurally hermaphrodite ones. Rudimentary stamens or pistils +are of very common occurrence in monstrous flowers. See Chapter on +Heterogamy, &c. + +FOOTNOTES: + +[465] 'Rev. Hortic.,' 1866, p. 467. + +[466] De Rochebrune, 'Bull. Soc. Bot. Fr.,' ix, p. 281. The author +points out seven grades between complete absence of petals and their +presence in the normal number in this plant. See also Gaudin, in 'Koch. +Fl. Helv.;' Koch. 'Synops. Fl. Germ.;' Cramer, 'Bildungsabweich,' p. 85. + +[467] 'Bull. Acad. Belg.,' t. xix, part 1, p. 255. + +[468] 'Bull. Bot.,' i, p. 7, tab. i, f. 7. + +[469] See Gay, 'Ann. Sc. Nat.,' iii, p. 27. + +[470] 'Ann. Sc. Nat.,' 4 ser., v, p. 305. + +[471] Cramer, 'Bildungsabweich,' p. 90. + +[472] See also Clos, 'Bull. Soc. Bot. Fr.,' xiii, p. 96, adnot. + +[473] See Cramer, 'Bildungsabweich,' p. 7. Hildebrand, 'Bot. Zeit.,' xx, +1862, p. 209. + +[474] See Hildebrand, 'Bot. Zeit.,' xx, 1862, p. 209. + +[475] 'Bull. Soc. Bot. Fr.,' viii, p. 287. + +[476] 'Bull. Soc. Bot. Fr.,' vol. viii, 1861, p. 152. + +[477] Ibid., ix, p. 275. + +[478] Ibid., 1861, vol. viii, p. 149. + +[479] 'Lobelia,' p. 55. + +[480] 'Phil. Bot.,' p. 119. + +[481] 'Cat. Plant. Pyr,' p. 58. + +[482] Moquin-Tandon, loc. cit., p. 328. + +[483] For other instances see Chatin in 'Ann. Sc. Nat.,' 4 ser., vol. v, +p. 305. + +[484] See also Morren. 'Bull. Acad. Belg.,' xv, Fuchsia, p. 67. + +[485] Cited in 'Bull. Soc. Bot., France,' t. xiv ("Rev. Bibl."), p. 253. + +[486] 'Primit. Flor. Amurens.' p. 57. + +[487] 'Flora.' 1848. p. 484. + + + + +BOOK IV. + +DEVIATIONS FROM THE ORDINARY SIZE AND CONSISTENCE OF ORGANS. + + +In the animal kingdom the entire adult organism, as well as each of its +separate parts, has certain dimensions, beyond which, under ordinary +circumstances, it does not pass, either in the one direction or the +other. It may not be easy or possible to state what the limits are, but, +practically, this inability to frame a precise limitation is productive +of no inconvenience. It is universally admitted that a certain animal +attains such and such dimensions, and that one organ has a certain +proportionate size as contrasted with another. The same rules hold good +in the case of plants, though in them it is vastly more difficult to +ascertain what may be called the normal dimensions or proportions. +Nevertheless observation and experience soon show what may be termed the +average size of each plant, and any disproportion between the several +organs is speedily detected. + +When there is a general reduction in size throughout all the organs of a +plant, or throughout all the nutritive organs, stem, leaves, &c., and +the several portions participate in this diminished size, we have what +are generally termed "dwarf varieties," dwarf in comparison, that is, +with the ordinary condition of the plants; on the other hand, if the +entire plant, or, at least, if the whole of one set of organs be +increased in size beyond the recognised average, we have large +varieties, often qualified by such terms as _macrophylla_, _longifolia_, +_macrantha_, &c. &c. In all these cases either the entire plant or whole +series of organs are alike increased or diminished beyond average +limits; and such variations are often very constant, and are transmitted +by hereditary transmission. It may be supposed that such deviations may +have originated, in the first instance, either from excessive use, or +from disuse, or from the agency of certain conditions promoting or +checking growth, as the case may be; but whether or no, it is certain +that these variations often persist under different conditions, and that +they often retain their distinctive characters side by side with plants +presenting the normal average dimensions. In other cases the variations +in size are of a less general character, and affect certain organs of a +whorl in a relative manner, as, for instance, in the case of didynamous +or tetradynamous stamens, where two or four stamens are longer than +their fellows, the long or short stamens and styles of di- and +tri-morphic flowers, &c. These differences are sometimes connected with +the development of parts in succession, and not simultaneously. + +Teratological deviations of size differ from those of which mention has +just been made chiefly in this, that they are more limited in their +manifestations. It is not, as a rule, the whole plant, or the whole +series of nutritive or of reproductive organs, that are affected, but it +is certain parts only; the alteration in size is more a relative change +than an absolute one. + +For convenience sake the teratological alterations of size may be +divided into those which are the result of increased growth and those +which arise from diminished action. It will be seen, therefore, that in +these instances it is the bulk of the organs that is increased, not +their number; moreover, their development or metamorphosis is not +necessarily altered. In connection with increased size an alteration of +consistence is so frequent that the two phenomena are here taken +together. It will be borne in mind that the changes of consistence from +membranous to succulent or woody are very frequent in the ordinary +course of development. They may also occur as accidental phenomena, or +the normal conditions of any particular flower or fruit may be exactly +reversed, the usually succulent fruit becoming dry and capsular, and so +forth. + + + + +PART I. + +HYPERTROPHY. + + +The term hypertrophy may serve as a general one to comprise all the +instances of excessive growth and increased size of organs, whether the +increase be general or in one direction merely. General hypertrophy is +more a variation than a deformity, unless indeed it be caused by insect +puncture or the presence of a fungus, in which case the excessive size +results from a diseased condition. For our present purpose hypertrophy +may be considered as it affects the axile or the foliar organs, and also +according to the way in which the increased size is manifested, as by +increased thickness or swelling--intumescence, or by augmented +length-elongation, by expansion or flattening, or, lastly, by the +formation of excrescences or outgrowths, which may be classed under the +head of luxuriance or enation. + +As size must be considered in this place relatively, it is not possible +to lay down any precise line separating what are considered to be the +normal dimensions from those which are abnormal. + +In practice no inconvenience will be found to accrue from this inability +to establish a fixed rule, and we may say that an hypertrophied organ is +one which, from some cause or other, attains dimensions which are not +habitual to the plant in its usual, healthy, well-formed state. + +It will be seen that under this general head of hypertrophy, increase of +size, however brought about, is included; thus, not only increase in +length, but also in thickness; alterations of substance or consistence, +no less than of dimensions, are here grouped together. The alterations +of consistence resulting from an inordinate development of cellular, +fibrous, or ligneous tissue, are, of course, strictly homologous with +the similar changes which occur, under ordinary circumstances, during +the ripening of fruits or otherwise. + +Hypertrophy, whatever form it may assume, may be so slight as not +perceptibly to interfere with the functions of the part affected, or it +may exist to such an extent as to impair the due exercise of its office. +It may affect any or all parts of the plant, and is generally coexistent +with, if not actually dependent on, some other malformation. Thus, the +inordinate growth of some parts is most generally attended by deficiency +in the size and number of others, as in the peripheral florets of +_Viburnum_ or _Hydrangea_, where the corollas are relatively very large, +and the stamens and pistils abortive. + + + + +CHAPTER I. + +ENLARGEMENT. + + +A swollen or thickened condition (_renflement_) is usually the result of +a disproportionate formation of the cellular tissue as contrasted with +the woody framework of the plant. We see marked instances of it in +cultivated carrots and turnips, the normal condition of the roots or +root-stocks in these plants being one of considerable hardness and +toughness, and their form slender, tapering, and more or less branched. + +The disproportionate development of cellular tissue is also seen in +tubers and bulbs, and in the swollen stems of such plants as +_Echinocactus_, _Adenium obesum_, some species of _Vitis_, &c. So, too, +the upper portion of the flower-stalk occasionally becomes much +dilated, so as ultimately to form a portion of the fruit. But it is not +necessary to give farther illustrations of this common tendency in some +organs to become hypertrophied. As a result of injury from insects or +fungi, galls and excrescences of various kinds are very common, but +their consideration lies beyond the scope of the present work. + +[Illustration: FIG. 200.--_Pelargonium_, one branch of which was +hypertrophied.] + +=Enlargement of axile organs.=--All the species of _Pelargonium_, +_Geranium_, _Mirabilis,_ as well as those of _Caryophylleae_ and other +orders, have tumid nodes as a normal occurrence. In the genus +_Pelargonium_ this swelling is sometimes not confined to the nodes, but +extends to the interspaces between them, _e.g._ _P. spinosum_. This +condition, which happens as a natural feature in the species just named, +may also occur as an exceptional thing in others. The author is indebted +to Dr. Sankey for a branch of _Pelargonium_ which was thus thickened, +the remaining branches not being in any way affected. The leaves on the +swollen branch were smaller than the others, and their stalks more +flattened. There was, in this instance, no trace of fungus or insect to +account for the swelling of a single branch, which might, therefore, be +due to bud-variation, perhaps to reversion to some ancestral form. The +repeated cross fertilisations to which Pelargoniums have been subjected +render this hypothesis not an improbable one. + +As an accompaniment to a spiral torsion of the woody fibres, this +distension of the stem is frequently met with, as in _Valeriana_, +_Dipsacus,_ &c. (See Spiral Torsion.) + +=Knaurs.=--On certain trees, such as the oak, the hornbeam, some species +of _Crataegus_, &c., hard woody lumps may occasionally be seen +projecting, varying greatly in size, from that of a pea to that of a +cocoa-nut. They are covered with bark, and consist in the interior of +very hard layers of wood disposed irregularly, so as to form objects of +beauty for cabinet-makers' purposes. From the frequent presence of small +atrophied leaf-buds on their surface, it would seem as if the structures +in question were shortened branches, in which the woody layers had +become inordinately developed, as if by compensation for the curtailment +in length.[488] The cause of their formation is not known, but it has +been ascertained that they are not due to insect agency. Knaurs may +occasionally be used for purposes of propagation, as in the case of the +"uovoli of the olive" and the "burrs" that are formed on some varieties +of apple, from which both roots and leaf-shoots are produced in +abundance. + +A distinction must be drawn between those instances in which the +swelling is solid throughout from the excessive formation of cellular +tissue, and those wherein it is hollow from the more rapid growth of the +outer as contrasted with the inner portions. These latter cases might be +classed under the head of distension. + +[Illustration: FIG. 201.--Formation of tubers or hypertrophied buds in +the axils of leaves in the potato.] + +=Enlargement of the buds= may be seen in the case of bulbs and tubers. +Occasionally these organs are developed in the axils of leaves, when +their nature becomes apparent. A swollen bud or bulbil in this +situation is not uncommon in some cultivated tulips and lilies. The +presence of small tubers in the axils of the leaves in the potato, as +shown in fig. 201, is also not unfrequent. + +[Illustration: FIG. 202.--Inflorescence of ash (_Fraxinus_), with +hypertrophied pedicels, flowers absent.] + +=Enlargement of the flower-stalk.=--The cauliflower and broccoli afford +familiar illustrations of hypertrophy of the flower-stalk, accompanied +by a corresponding defective development of the flowers. In the case of +the ash the terminal pedicels occasionally become swollen and distorted, +while the flowers are completely deficient, as shown in the adjacent cut +(fig. 202). + +In grapes a similar condition may occasionally be met with in which the +terminal pedicels become greatly swollen and fused into a solid mass. It +would seem probable that this change is due to insect puncture, or to +the effect of fungus growth at an early stage of development, but as to +this point there is at present no evidence.[489] + +[Illustration: FIG. 203.--Monstrous pear, showing extension and +ramification of the succulent floral axis. The bases of the sepals are +also succulent.] + +In the apple a dilatation of the flower-stalk below the ordinary fruit +may occasionally be observed, thus giving rise to the appearance of two +fruits superposed and separated one from the other by a constriction. +(See fig. 176, p. 327.) The lower swelling is entirely axial in these +cases, as no trace of carpels is to be seen. M. Carriere[490] mentions +an instance wherein from the base of one apple projected a second +smaller one, destitute of carpels, but surmounted by calyx-lobes as +usual. The direction of this supernumerary apple was the exact opposite +of that of the primary fruit. + +[Illustration: FIG. 204.--Monstrous pear, showing extension and swelling +of axis, &c.] + +In pears, quinces, and apples, a not uncommon deviation is one in which +the axis is prolonged beyond the ordinary fruit, like which it is much +swollen. Occasionally the axis is not only prolonged, but even ramifies, +the branches partaking of the succulent character of the ordinary pome. +Such instances are frequently classed under the head of prolification, +but they have in general no claim to be considered in this light, for +the reasons already given in the chapter relating to that subject. (See +p. 135.)[491] + +A very curious illustration of hypertrophy of the flower-stalk is +recorded and figured by M. Carriere[492] in the cherry. The calyx in +these fruits was completely superior, the succulent portion of the fruit +being made up of the dilated extremity of the peduncle, and possibly in +part of the base of the calyx. The general appearance was thus that of a +crab-apple. There was no stone in the interior, but simply a rudimentary +kernel or seed.[493] + +Moquin-Tandon records an instance in which the stamens of each +individual flower in the inflorescence of a vine were hypertrophied, the +sepals, petals, and other organs of the flower, being proportionately +diminished.[494] + +In this place may also be mentioned the hypertrophied condition of the +placenta observed by Alphonse de Candolle in a species of _Solanum_, and +also in a species of _Melastoma_. Not only was the placenta unusually +large in these flowers, but it also protruded beyond the ovary.[495] A +similar state of things in _Lobelia_ and _Cuphea_ has already been +alluded to under the head of Alterations of Direction (p. 210). + +The following singular growth in a tomato is described by the Rev. M. +J. Berkeley in the 'Gardeners' Chronicle' for 1866, p. 1217, and appears +to have been an extension of the placenta:--"On the first glance it +seemed as if an unusually large grape-stone had accidentally fallen on +the upper surface of the fruit, and was attached by the narrow base. The +process was, however, five lines long, and much narrowed below, besides +which, though it was pale green above, the base was coral-red, like the +tomato itself. It grew on a narrow and shallow crack on the surface of +the fruit, and was found below to communicate directly with a +fibro-vascular bundle, which entered into the composition of a portion +of the placenta. On making a vertical section, instead of being +succulent, as I expected, it was white and spongy within, with several +lacunae, and one or two irregular fibro-vascular bundles, with highly +developed spiral vessels threading the centre. These vessels, moreover, +were tinged with brown, as in many cases of diseased tissues. There was +not the slightest appearance of placentae or anything indicating an +abortive fruit. On closer examination the cuticle was found to consist +of thick-walled cells, exactly like those of the tomato, while the +spongy mass consisted of a similar tissue to the fleshy portion of the +fruit, but with far less wrinkled walls, and more indistinct +intercellular spaces. The most striking point, however, was the immense +quantity of very irregular and unequal starch-grains with which they +were gorged, which gave a peculiar sparkling appearance to them when +seen _en masse_. I am inclined to regard the body rather as an abortive +axis than an undeveloped fruit. In almost all, if not all, these cases +of abnormal growth, whether from leaves, petioles, fruit, or other +portions of the plant, we find an immediate connection with one or more +spiral vessels, which if not existent at first are developed sooner or +later. In the present case the connection of the fibro-vascular tissue +of the fruit and abnormal growth was plain enough, but whether it +existed when the body was first given off I am unable to say, as it was +fully developed when the fruit was brought to me." + +=Enlargement of the leaves.=--Increase in the size or substance of +leaves takes places in several ways, and affects the whole or only +certain portions of them. The simplest form of this malformation is met +with in our cabbages, which, by the art of the gardener, have been made +to produce leaves of greater size and thickness than those which are +developed in the wild form. In such instances the whole substance of the +leaf is increased in bulk, and the increase affects the fibrous +framework of the leaves as well as the cellular portions, though the +exaggerated development of the latter is out of proportion to that of +the former. + +In some species of _Podocarpus_ there may occasionally be seen at the +base of the branchlets a dozen or more fleshy scales, of a rose colour, +passing gradually into the ordinary leaves of the plant, and evidently +analogous to the three fleshy confluent bracts which surround the ripe +fruit. + +In other instances, while the fibrous framework of the leaf retains its +usual degree of development, the cellular parenchyma is developed in +excess, and, if the increase is so arranged that the number of +superposed layers of the cellular tissue is not increased, or their +thickness exaggerated, then we get such leaves as those of the "kail," +or of the "Savoys" leaves, which are technically called by descriptive +botanists "folia bullata." In such leaves the disc of the leaf, rather +than the margin, is increased and its surface is thrown up into little +conical projections, which are hollow on the under side. + +But leaves may increase beyond their usual size without such grave +alterations of form as those to which allusion has just been made. It is +well known that if a tree be cut down and new shoots be sent out from +the stump, the leaves formed on these shoots very often greatly exceed +the ordinary ones in dimensions. Such cases as this hardly come under +the head of malformations. But where one part only of the leaf is +excessively developed, the other portion remaining in its ordinary +condition, there can be no hesitation in ranking the phenomenon as +teratological. + +Thus, Moquin says that the median nerve may be prolonged beyond the +blade of the leaf in the form of a short strap or ribbon-like +excrescence, while, at other times, the lateral parts of the leaf are +subjected to undue development. He refers to a case cited by +Schlotterbec[496] in which each side of the leaves of a yellow "violier" +(wallflower) was dilated into a kind of projecting lobe on either side +of the true apex of the leaf, thus rendering it in appearance +three-lobed. M. Delavaud[497] puts on record a case of hypertrophy in +the leaves of the common elm, resulting in the formation of an +additional lobe and a return to the tricostate type. A leaf so affected +is stated to have presented the appearance of a fusion of two leaves. +(See also Multiplication of leaves, p. 353.) + +The hypertrophied and coloured leaf of _Gesnera_ occupying the place of +the absent inflorescence has been previously alluded to under the head +of displacement (p. 88). + +In some instances hypertrophy is the opposite of suppression; as in the +case previously mentioned, where the stipule in the inflorescence of a +pea, which is usually undeveloped and rudimentary, was developed in the +form of a leafy cup or pitcher. + +Another instance of the development of parts usually suppressed, is +afforded by the bud-scales of _Magnolia fuscata_, which may sometimes be +found with small but perfect leaves projecting from them, the leaf in +this case being the lamina which is ordinarily abortive, while the +scales are the representatives of the stipules. This condition is said +by Hooker and Thomson ('Flora Indica,' p. 73) to be constant in +_Magnolia Campbelli_. + +=Enlargement of the perianth, &c.=--One or all the segments of the +perianth may be subjected to hypertrophy; thus, the utricle of _Carex +vulpina_ may frequently be observed to attain four or five times its +usual size, the contained ovary remaining unaffected. This condition is +generally the result of insect puncture. The growth of parasitic fungi +will produce a similar result, as is often seen in the common shepherd's +purse, _Thlaspi bursa pastoris_, and other _Cruciferae_. The perianth of +_Rumex aquaticus_ has been also observed to be occasionally +hypertrophied in conjunction with a similar condition of the pistil and +with atrophy of the ovules. + +Moquin relates having found flowers of _Salsola Kali_ and of +_Chenopodium murale_ in which some of the segments of the perianth were +five or six times larger than they should be. + +[Illustration: FIG. 205.--Hypertrophy of the perianth in _Cocos +nucifera_.] + +The adjoining woodcut represents a singular condition of some cocoa-nuts +in the Kew Museum, the appearance of which is due apparently to an +hypertrophied condition of the segments of the perianth, which have not +only increased in length as the central nut has ripened, but have +developed in their tissues that fibrous tissue which ordinarily is found +in the pericarp only. This view of the structure of these nuts is borne +out by the fact that, under normal circumstances, the base of the +perianth contains a considerable amount of fibrous material. In the +present case this has increased to such an extent that the fruit appears +surrounded by a double husk, by an inner one as usual, and by an outer +six-parted one. + +It will be remembered that in some of the _Cinchonaceae_, e.g. +_Mussaenda_, _Pinckneya_, _Calycophyllum_, one or more of the calycine +lobes are normally dilated and petaloid, the others remaining small and +comparatively inconspicuous. Inequality in size is, indeed, a common +occurrence in the sepals of many natural orders--_Polygalaceae_, +_Leguminosae_, _Labiatae_, &c. The flowers of a rose are mentioned by +Moquin as having presented an enlargement of the calyx without any other +alterations in form. Schlechtendal has noticed the same thing in +_Papaver Rhoeas_, Reichenbach in _Campanula persicifolia_, and A. de +Candolle in _C. Rapunculus_. M. Brongniart also has recorded[498] a +remarkable variety of _Primula sinensis_ cultivated in the Jardin des +Plantes at Paris, wherein the calyx is enormously developed. MM. +Fournier and Bonnet have described flowers of _Rubus_ with hypertrophied +calyx in conjunction with atrophy and virescence of the petals and other +changes.[499] + +The corolla may be hypertrophied in some cases, though the change is +more rare than in most other organs. Moquin-Tandon mentions as subject +to this anomaly species of _Galeopsis_, _Prunella_, _Scabiosa_, and +_Dipsacus_, and also mentions a remarkable variety of _Viola odorata_ +cultivated in the neighbourhood of Toulouse. The same learned author +also alludes to the so-called double Composites, viz. those in which the +usually tubular florets of the disc assume the form and proportions of +those of the ray, but these are hardly cases of hypertrophy. + +=Enlargement of the androecium.=--Dunal[500] alludes to a curious +instance in a species of _Verbascum_, the lower flowers of which had +hairy stamens as usual, but the filaments of the topmost flower were +quite destitute of hairs, and dilated like a flat ribbon. + +Moquin relates having found in the neighbourhood of Toulouse a plant of +_Solanum Dulcamara_ in which all the upper flowers had two or three +stamens of larger dimensions than the others. This happens habitually in +_Solanum tridynamum_ and _S. Amazonicum_, and to a less extent in _S. +vespertilio_ and _S. cornutum_; also in some species of _Hyoscyamus_. +These cases show the close affinity between the _Solanaceae_ and the +_Scrophulariaceae_. + +=Enlargement of the gynoecium.=--In some flowers which have become +accidentally female the pistil becomes unusually large, and even to such +an extent as to prevent the passage of the pollen. Moquin remarks having +seen this enlargement in the pistils of _Suaeda fruticosa_ and _Kochia +scoparia_. The flowers of these Chenopods, under these circumstances, +resemble the female flowers of some nettles. The styles of _Anemone_ are +also much enlarged as the result of cultivation, and from their petaloid +appearance resemble those of the _Iris_ (Goethe). MM. Seringe and +Heyland[501] have figured some anomalous flowers of _Diplotaxis +tenuifolia_ in which the pistil, more or less distended and deformed, +was considerably elongated below, so that it seemed to be borne upon a +long stalk, analogous to that of fruits of Capparids. Dr. Klinsman[502] +mentions an instance of a similar kind combined with hypertrophy of the +sepals and pistils; indeed, the alteration is not uncommon among +Crucifers. _Pyrethrum inodorum_ is very subject to hypertrophy. The +styles of its radial florets become elongated without any other +alteration; at the same time the small corollas become green, and show a +tendency to assume a foliaceous condition. Sometimes the hypertrophy +affects also the styles of the central florets, and these also become +enlarged to double or treble their usual dimensions. + +Linne has remarked that the ovary of _Tragopogon_ sometimes assumes very +large dimensions, as also does the pappus. He mentions a double-flowered +variety, the ovaries of which become ten or twelve times larger than +ordinary. M. Clos[503] records an instance in _Rumex scutatus_ wherein +the pistil was hypertrophied or club-shaped, and open at the top, or in +other cases funnel-shaped, three-lobed at the summit, each lobe +terminated by a style. One of the most frequent causes tending to the +hypertrophy of the pistil is attributable to the puncture of insects; +thus, when the ovary of _Juncus articulatus_ is thus punctured, it +acquires a size two or three times larger than ordinary, becoming at the +same time sterile.[504] + +Occasionally the enlargement may be due to a fusion or incorporation of +other elements; thus, M. Lemaire describes an instance in which the +style of _Sinningia purpurea_ was much larger than ordinary, tubular, +bearing three small lobes, and altogether bearing much resemblance to +the column or "gynosteme" of Orchids. This appearance was due to the +cohesion and intimate union of the styles with three abortive +stamens.[505] + +=Enlargement of the fruit.=--Most cultivated fruits are in a state of +true hypertrophy. Girod de Chantrans, after many trials, succeeded in +producing a peculiar variety of pea with pods double the ordinary +size.[506] M. Clos[507] mentions a case wherein the carpels of +_Delphinium dictyocarpum_ were hypertrophied. The change in size may or +may not be attended by a difference in form; thus, in certain +_Leguminosae_, as _Medicago lupulina_, _Melilotus leucantha_, the carpels +are sometimes hypertrophied and elongated, so as to resemble a claw or +hook.[508] + +The fruit of the common groundsel (_Senecio vulgaris_) is in its normal +condition two or three times shorter than the involucre, and cylindrical +for its whole length, but it frequently happens that the fruits become +as long as the involucre itself, and taper from the base upwards, so as +to become beaked. Under this head may also be mentioned the fleshy +bulbils that are found in the capsules of _Crinum_, _Amaryllis_, and +_Agave_. These are true seeds enormously dilated.[509] In these seeds +the outer coating becomes very thick and fleshy, and is traversed by +spiral vessels. + +It is obvious that very important results in a practical point of view +may be and have been arrived at by cultivators availing themselves of +this tendency of plants to increase in dimensions under certain +circumstances. It is needless to do more than refer to the many fruits, +vegetables, and cereals, which have thus become enlarged and improved by +careful selection and rearing. + +=Alterations of consistence= often accompany changes in size. The change +may be one whereby the tissues become unusually hardened, by the +excessive formation of secondary woody deposits, or softer and more +succulent than ordinary, from the formation of an inordinate amount of +loose cellular tissue. Generally speaking, the appearances presented in +such cases are not sufficiently striking to demand notice other than as +regards their size. One illustration, however, may be cited from its +singularity. This was the case of a dahlia, in which the centre of the +flower was occupied by a projecting knob as large as a walnut, brown in +colour, and very hard in texture. This knob was nothing but the enlarged +and indurated extremity of the common receptacle, destitute of the +scales and florets which usually spring from it. No insect-puncture +could be detected, and no other reason for this peculiarity could be +ascertained. + +FOOTNOTES: + +[488] On the subject of knaurs, the reader is referred to Trecul, 'Ann. +Sc. Nat.,' 3 ser., vol. xx, p. 65; Lindley, 'Theory of Horticulture;' +Rev. M. J. Berkeley, 'Gardeners' Chronicle,' 1855, p. 756. + +[489] Jaeger, 'Flora.' 1860. p. 49, tab. i. + +[490] 'Revue Horticole,' 1868, p. 110, figs. 12, 13. + +[491] The reader may also refer for further information on the subject +of malformed pears to Irmisch. 'Flora,' 1858, p. 38, tab. i; Lindley, +'Theory of Horticulture'; Caspary, 'Bull. Soc. Bot. France,' vol. vi, +1859 (Rev. Bibl.), p. 235; Duhamel, 'Phys. Arbr.,' liv. iii, cap. 3. p. +393, fig. 308; Bonnet, 'Recherch. Us. feuilles,' tab. xxvi, fig. 2; +Moquin-Tandon, 'El. Ter. Veg.,' p. 384, &c. Some of the cases recorded +are, however, instances of true prolification. + +[492] 'Revue Horticole' 1868, p. 310. + +[493] The interest of this accident is great, as showing how an +habitually superior ovary may become inferior--a change so rare in its +occurrence that its existence has been denied, and thus forming a marked +contrast with the frequency with which the converse change of an +inferior ovary to a superior one, from want of union with the calyx or +from imperfect development of the peduncle, may be observed. It is also +interesting as showing how the peduncle may become swollen, and at the +same time how the woody deposit of the endocarp may, as if by +compensation, be deficient. And, again, the malformation is not without +significance in regard to the relationship between the drupaceous and +the pomaceous subdivisions of _Rosaceae_. The case would fitly be +included under alterations of position, but the sheets relating to that +subject were printed off before the publication of M. Carriere's notice. + +[494] 'Bull. Soc. Bot. France,' 1860, vol. vii, p. 881. + +[495] "Monstr. Veget.," in 'Neue Denkschrift.' + +[496] "Sched. de Monst. Plant." in 'Act. Helvet.,' t. ii, pl. ii, f. 14. + +[497] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 144. + +[498] 'Ann. Sc. Nat.,' ser. 2, t. i, p. 308, pl. ix _c_, fig. 1. + +[499] 'Bull. Soc. Bot. France,' 1862, t. ix, p. 37. + +[500] 'Consid. org. Fleur.,' Montpell., 1829, 25, 26, pl. ii, f. 18 and +19. + +[501] 'Bull. Bot.,' t. i, p. 7, tab. 1. + +[502] 'Linnaea,' vol. x, p. 604, tab. 5. + +[503] 'Mem. Acad. Sc. Toulouse,' 5 ser., vol. iii. + +[504] 'Re. nosol. Veget.,' pp. 342. + +[505] 'Illustr. Hortic.,' 1868, Misc., p. 62. + +[506] 'Ann. Soc. Linn.,' Paris, t. i, p. 139. + +[507] 'Mem. Acad. Toulouse,' t. 6, 1862. + +[508] 'D. C. Prod.,' ii, pp. 172, 187. + +[509] Richard, "Obs. sur les bulbilles des Crinum;" 'Ann. Sc. Nat.,' t. +ii, p. 12. pl. i, fig. 1, 2. See also A. Braun, "Memoire sur les graines +charnues des Amaryllidees," &c.; 'Ann. Sc. Nat.,' 1860, vol. xiv, p. 1, +tab. 1. + + + + +CHAPTER II. + +ELONGATION. + + +The class of cases coming under this head are sufficiently indicated by +the name. There are many instances of this phenomenon occurring under +different conditions, which, though unusual, can hardly be called +abnormal, such, for instance, as the great lengthening of roots in their +search for water, the excessive elongation that takes place in plants +when grown at a distance from the light, in their endeavour to attain to +which they become, as gardeners phrase it, "drawn." A similar result is +brought about in forests or plantations, where long spars are required, +by allowing the trees to grow very close to each other, so as to prevent +the lateral extension of the branches. When plants grow in running +water their roots, stems, and sometimes their leaves, become excessively +elongated, as in _Ranunculus fluitans_, the flower-stalks of _Valisneria +spiralis_, &c. These are cases of variation rather than of malformation, +but are none the less curious, or sometimes perplexing; thus, Lapeyrouse +described, in his 'Supplement a la flore des Pyrenees,' p. 27, under +the name _Potamogeton bifolium_, a plant which Mr. Bentham subsequently +discovered to be nothing but a flowerless variety of _Vicia Faba_ +distorted by its growth in water.[510] + +=Elongation of the root.=--This, as already remarked, is more often a +variation than a malformation, and is usually due to the presence of +water at a distance necessitating growth at the extremities of the root, +or to the presence of some obstacle, such as a stone, to avoid which the +root elongates till it has passed the obstruction. Occasionally in +Crocus corms some of the fibrils may be met with much lengthened and +thickened, and invested with a fleshy sheath. It is not certain, +however, that these structures are roots; possibly, nay probably, they +may be processes from the stem thrust downwards into the soil, similar +to the formations already described in the tulip (p. 85, fig. 39). + +=Elongation of the inflorescence.=--Under this heading it is necessary +to consider lengthening of the common rachis in the case of an aggregate +inflorescence, and lengthening of the individual flower-stalks, whether +they be solitary or portions of a multiple inflorescence. The two +phenomena may occur together, but they are quite as often independent +one of the other. Thus, among _Umbelliferae_ the umbels are occasionally +met with supported on unusually long stalks, while the pedicels of the +individual flowers may or not be increased in length; so also with some +of the Composites, or the heads of flowers of some _Leguminosae_, +_Trifolium repens_, &c. &c. + +Another illustration of the sort is that recorded by M. Fournier, +wherein the usually umbellate inflorescence of _Pelargonium_ was, +through the lengthening of the main stalk, transformed into a raceme. +Among Composites a similar change may sometimes be met with. + +MM. Clos and De Schoenefeld have recorded the existence of a variety of +the sweet chestnut (_Castanea_)in which the female catkins were as long, +and bore nearly as many flowers, as the male spikes. This is stated to +be of constant occurrence in some localities, and to be accompanied by a +diminished size of the fruits. A similar elongation has been observed in +the case of the walnut, catkins of which have been seen bearing thirty +to thirty-five large nuts.[511] + +In the strobile of the hop, _Humulus Lupulus_, a like elongation may +sometimes be met with, generally in association with a more or less +leafy condition of some of the scales. + +Of a similar character, but complicated with extrusion or eversion of an +ordinarily concave axis, is the fig described by Zuccarini,[512] and +from the appearances presented by which that author draws the inference +that the peculiar appearance of the fig is due to the formation of a +large number of small bracts blended together for the greater part of +their length, and accompanied by the suppression of the internodes, and +consequent shortening of the axis. In the monstrous fruit the axis is +prolonged, and forms a kind of raceme or catkin, surrounded at the base +by numerous bracts, as in many _Amentaceae_. (See p. 204, figs. 105, +106.) + +A lengthening of the axis of the female strobiles of _Coniferae_ is not +of infrequent occurrence in _Cryptomeria japonica_, _Larie europaea_, +&c., and this is usually associated, as has been before stated, with a +leaf-like condition of the bracts, and sometimes even with the +development of leaf-bearing shoots in place of the scales. (See under +Prolification of Inflorescence and Phyllomorphy, and for references, p. +115.) + +=Elongation of the secondary flower-stalks.=--In the previous section +the effect of elongation of the main rachis has been considered. A +corresponding deviation occurs in the peduncles or pedicels, and +sometimes alters the general character of the inflorescence very +considerably, converting a spike into a raceme, a raceme into a corymb, +a capitulum into an umbel, and so forth. A few such cases may here be +alluded to. Fig. 206 represents a specimen of _Ranunculus acris_, in +which the lower and lateral flower-stalks were not only increased in +number, but so much lengthened as to form a flat-topped inflorescence--a +corymbose cyme. In many leguminous plants, as in _Trifolium repens_, +_Lotus corniculatus_, &c., what is usually a compact spike, or head of +flowers, becomes a raceme from the elongation of the pedicels. In +_Umbelliferae_ a similar change occurs, by virtue of which sometimes the +umbels themselves, and at other times the florets, are raised on +unusually long stalks, as in _Angelica Razoulzii_, _Carum Carui_, +_Thysselinum palustre_.[513] In _Compositae_, when affected by an +analogous change, the capitulum assumes the appearance of a simple +umbel, as in _Hypochaeris radicata_, _Senecio vulgaris_, and other +plants. + +[Illustration: FIG. 206.--Inflorescence of _Ranunculus acris_, with +secondary peduncles lengthened.] + +In some of the double-flowered apples which have been previously alluded +to, the flower-stalk is inordinately long when compared with the +adjacent ones. Possibly in some of these cases the absence of the usual +swelling of the upper part of the peduncle may be connected with its +increased length. One of the most striking instances of lengthened +flower-stalk occurred in an apple flower, wherein there was no swelling +beneath the calyx, while the latter was represented by five perfect +stalked leaves. + +=Elongation of the leaves.=--In the case of water plants this change +keeps pace with the corresponding growth of the stem, _e.g._ +_Ranunculus fluitans_, and in terrestrial plants there are varieties +termed longifoliar, from the unusual length of the leaves. A similar +lengthening occurs in the involucral leaves of _Umbelliferae_ and +_Compositae_, changing very materially the general aspect of the +inflorescence. Occasionally, also, the leaf-lobes of parsley (_Apium +Petroselinum_) and other crested-leaved plants may be observed to lose +their ordinary wavy form, and to be lengthened into flat riband-like +segments, as shown in fig. 207. + +The only further illustrations that it is requisite to give of such +changes in this place are those occurring in lobed or compounded leaves, +which, from a lengthening of the midrib or central stalk, convert a +digitate or palmate leaf into a pinnate one. In these instances the +lobes or leaflets become separated one from another by a kind of +apostasis. This change may be frequently seen in the horse-chestnut, +particularly in the young shoots formed after the trees have been pruned +or pollarded. In the adjoining cut the intermediate stages between a +palmate or digitate leaf to a pinnate one may be seen. The specimens +from which the drawing was made were taken from the same tree at the +same time. + +[Illustration: FIG. 207.--Portion of leaf of parsley, showing the change +from short wavy to long flat leaf-lobes.] + +In the white clover, _Trifolium repens_, a similar transition may often +be observed, as also in some species of _Potentilla_.[514] + +=Elongation of the parts of the flower.=--The only circumstance that +needs especial mention under this section is the great lengthening that +sometimes takes place in the carpels, sometimes as a result of injury +from insects or fungus, at other times without assignable cause. + +[Illustration: FIG. 208.--Leaves of horse-chestnut, _AEsculus_, showing +passage from digitate to pinnate leaves.] + +In the case of inferior ovaries this lengthening is, perhaps, even more +common, as in _Umbelliferae_, _Compositae_, &c. The common groundsel +(_Senecio vulgaris_) is especially liable to this form of enlargement of +the pistil, either in association with a leafy condition of the pappus +or without any such change. + +=Elongation of the thalamus, placenta, &c.=--In some plants, as in +_Magnolia_ or _Myosurus_, the thalamus becomes much elongated, and bears +the carpels disposed spirally around it. A similar lengthening occurs in +malformed flowers, usually in association with a similar change in the +lower or outer part of the flower, by virtue of which the whorls become +separated from each other (Apostasis). Elongation and protrusion of the +placenta have been already alluded to at p. 119, and also at p. 125. In +some of these cases the elongated placenta has taken the form of a +leaf-bearing shoot.[515] + +=Apostasis.=--Engelmann made use of this term to express the separation +of parts one from another by the unusual elongation of the +internodes.[516] He drew a distinction between the separation of +individual organs one from the other, and the corresponding displacement +of whorls. The subject has already been, to a considerable degree, +treated of in these pages under the head of dialysis, displacement, and +prolification, and but little need here be added. With reference to the +distance between one whorl and another, it will be remembered that, +although in the majority of cases the floral whorls are packed closely +together, yet in other instances the floral axis becomes elongated, and +thus separates the whorls one from another, by structures such as the +gynophores, androphores, &c., of _Passifloreae_, _Caryophylleae_, +_Capparideae_, &c. &c. + +A similar elongation of the thalamus, bringing about the separation of +the floral whorls, or of their constituent parts, is very commonly met +with in association with median prolification. Where the individual +floral elements are thus thrown out of their usual verticillate +arrangement, they naturally assume a spiral disposition, and are, in +some cases, united by their margins, so that a spiral sheet or tube is +formed, surrounding the axis. This frequently occurs in double flowers +of the Chinese primrose, _Primula sinensis_. + +Engelmann[517] figures a case wherein the calyx of _Anagallis +phoenicea_ was separated by a rather long internode from the corolla, +and a like illustration in _Torilis Anthriscus_. + +[Illustration: FIG. 209.--Flower of _Delphinium_, showing apostasis of +carpels, from lengthening of the thalamus, &c. (Cramer.)] + +A frequent change in Crucifers is due to the formation of a long stalk +bearing the pod, and thus giving rise to the appearance met as a +constant occurrence in Capparids. + +In _Tropaeolum majus_ a similar elevation of the pistil may occasionally +be seen. + +The adjacent figure of a monstrous _Delphinium_ taken from Cramer +illustrates well the elongation of the floral axis and the apostasis of +the carpels. In this instance the axis is terminated by a second flower +(median prolification). + +One of the best-marked illustrations of these changes occurs in a +permanent malformation of _Epilobium hirsutum_, specimens of which were +originally obtained from the late Professor Henslow. The several floral +parts are here, some virescent, others truly foliaceous, and each whorl +is separated from its neighbour by a rather long internode. In _Fuchsia_ +and _Campanula_ a like change may occasionally be observed. + +Engelmann, in addition to those previously mentioned, cites the +following plants as having manifested this change: + +_*Convallaria majalis!_, _*Tulipa Gesneriana!_, _Veronica Chamaedrys_, +_Orobanche gracilis_, _Solanum Lycopersicum_, _Gentiana campestris_, +_Hypericum_, _Helleborus fetidus_, _Caltha palustris_, _Brassica +oleracea!_ and many _Rosaceae_, _Caryophylleae_, _Cruciferae_, and +_Ranunculaceae_. (See Dialysis, Median Prolification, &c.) + +Apostasis of the sub-floral or involucral leaves is not of infrequent +occurrence in malformations affecting _Compositae_ and _Umbelliferae_. In +the following genera it has been observed with especial +frequency:--_Torilis Anthriscus_, _Eryngium_, _Athamanta Cervaria_, +_Leontodon_, _Tragopogon pratense!_, _Wedelia perfoliata!_ In garden +anemones, also, it is a common deviation. + +FOOTNOTES: + +[510] 'Cat. Plant.,' Lang., p. 113. + +[511] 'Bull. Soc. Bot. France,' t. i, 1854, p. 173, and t. xiii, p. 96. + +[512] 'Abhandl. Math. Phys. Class.,' Band. iv, Abhandl. i, tab. i. + +[513] See Cramer, 'Bildungsabweich,' pp. 62-79, and Fleischer, +'Missbild, der Culturpflanzen.' + +[514] Schlechtendal, 'Bot. Zeit.,' 1844, p. 457; 'Linnaea,' xi, p. 301, +xiv, p. 363; 'Bot. Zeit.,' 1856, p. 72; Masters, 'Rep. Brit. Assoc.,' +Manchester, 1861; Coultas, 'What may be learnt from a tree,' p. 118. + +[515] For further details refer to the chapter on Displacements, p. 86. + +[516] 'De Antholysi,' p. 42, Sec. 49. + +[517] Loc. cit., tab. 2, f. 6. + + + + +CHAPTER III. + +ENATION. + + +Under the above heading are included certain forms arising from excess +not of growth, but of development, and consisting in the formation of +supplementary lobes or excrescences from various organs. + +The new formations are not due either to a repetition or to a partition +of any organ, but are out-growths from others previously formed. + +In prolification and in multiplication the adventitious structures are +of independent origin. In fission the new developments grow +simultaneously with the older ones, of which, indeed, they are mere +repetitions. Moreover, in fission the supplementary lobes do not, in +general, project a plan different from that of the original structure, +at least in the first instance, though their direction may ultimately +become changed. + +In enation the new growth projects from a previously formed organ after +it has attained to considerable size, or even after its ordinary +proportions have been attained, and it sprouts out from the beginning in +a plane which is at a considerable angle to that of the parent organ, +and it is sometimes of a different structure from it, and has different +functions to fulfil. + +Many of the instances that occur of scales projecting from petals, as in +_Caryophylleae_, _Sapindaceae_, &c., the coronal filaments of +passion-flowers, the cup of _Narcissus_, the appendages that beset the +segments of the perianth in _Lilium lancifolium_, and other similar +growths, may be referred to a like process. In many cases this has been +proved by a study of the development of the flower, from which it +appears that the growths in question are developed subsequently to the +formation of the ordinary floral whorls. It is requisite, however, to be +cautious in pronouncing upon the exact nature of these bodies, in the +absence of a knowledge of their period and mode of formation. They may +be mere outgrowths from one or other of the customary whorls, or they +may represent abortive stamens or petals, &c. Where circumstances +prevent the course of development from being traced, something may be +inferred as to their real nature from their position in regard to the +other parts of the flower, from their anatomical structure, and from +analogy or comparison with like organs in other plants. The period of +their formation is, perhaps, of less importance than was at one time +supposed, since it is well ascertained that, in some cases, the +formation of the parts of the flower, _e.g._ the stamens of mallows, +follows a centrifugal rather than a centripetal order. + +In the case of monstrous developments of this nature too much care can +hardly be exercised, and the observer should rarely venture on an +explanation of the nature of the case from the evidence afforded by the +monstrous growth apart from that to be derived from the study of the +development and organization of the normal flower and from analogous +formations in allied plants. + +=Excrescences from axile organs--Warts.=--In a preceding paragraph the +formation of gnaurs has been alluded to. There are other outgrowths, +called warts, occasionally met with in trees, and which are more closely +connected with the central tissues of the stem, while at the same time +they are not provided with buds, in which two particulars warts differ +from gnaurs. + +Excrescences of this kind often attain a very large size, and may be +seen on old elms and other trees, but, as their formation is probably +more pathological than teratological, no further notice of these +structures need here be given. No special notice need here be taken of +the tubercles on the roots of so many _Leguminosae_, nor of the peculiar +excrescences on the roots of _Taxodium distichum_, as these appear to be +normal formations. But it may be well to mention in this place an +anomalous development which occurs occasionally in _Ruscus aculeatus_, +and in which, from the upper surface of the ordinary flattened leaf-like +branch, projects at right angles a second similar branch, so that in +section the appearance would be like that of the inverted letter t; +thus, [Symbol: Inverted upper-case T]. + +=Enation from foliar organs--Leaves.=--The development of adventitious +lobes from leaves may take place either from their surfaces or their +margins. A few illustrations may be given of each. In cabbage leaves a +formation of adventitious laminae projecting at right angles from the +primary one may frequently be observed. In the instance figured (fig. +210) the new growths proceeded almost exclusively from the thick midrib, +which, in the figure, is shown cut through just above the base. Not only +is the ordinary semilunar band of vascular tissue to be here seen, but a +similar broken line of vessels exists on the upper side of the +leaf-stalk; thus the whole structure resembles that of a stem or a +branch as much as that of a true leaf. + +[Illustration: FIG. 210.--Section through base of midrib of cabbage +leaf, showing supplementary laminae, &c.] + +The development of secondary leaves from the surfaces of primary ones +(phyllomania, autophyllogeny) has already been alluded to at p. 355. + +Some of the cases wherein a leaf seems to have a double lamina may be +alluded to here, though possibly they would more properly be referred to +fission. The appearance presented is as if four wings projected from the +midrib, so that a cross section would be nearly in the form of +[Symbol: )O( turned 90 degrees.]. In an orange leaf presenting this +appearance the lower surface of one lamina was, as usual, dull in +colour, while the upper surface of the subjacent lamina was likewise +dull; hence the impression might arise that this was an instance of the +adhesion of two leaves back to back, but the petioles were not twisted, +as they must have been had two leaves thus been united, and neither in +the petiole nor in the midrib was there the slightest indication of +fusion, the vascular bundles being arranged in a circular manner, not in +a horseshoe-like arrangement, as would have been the case had adhesion +taken place.[518] (See p. 33.) + +Such leaves as those of the hedgehog holly, _Ilex Aquifolium_, var. +_feroae_, and, to a less extent, bullate leaves, may also be mentioned +here as illustrations of hypertrophy or enation. + +[Illustration: FIG. 211.--_Nephrodium molle_. Ordinary frond and forked +and crested varieties of the same, the crest arising from the inordinate +development of the margins of the pinnules.] + +When the increased development occurs at the margin of the leaves, +especially, the result is a wavy or crisped appearance, "folia undulata, +_vel_ crispa."[519] These conditions occur normally in such leaves as +those of _Rumex crispus_, _Malva crispa_, &c., and are developed to an +extreme degree in garden varieties of parsley, some kails, &c., as well +as in many ferns, but these are probably cases rather of fission than +enation as here understood.[520] + +=Enation from the sepals.=--The basal lobes of the calyx in _Campanula +Medium_, under normal circumstances, may be referred to in illustration +of this occurrence, while the adventitious spurs on the calyx of some +monstrous flowers seem due also to a like cause. These have already been +alluded to at p. 315. + +=Enation from the corolla.=--The instances of this are more frequent +than in the case of the calyx, and admit of classification according as +they occur in polypetalous or gamopetalous flowers, on the outer or +inner surface of the petals, &c. Under natural circumstances the +formation of scales, lobes, &c., from the petals, as in some +_Caryophylleae_, _Sapindaceae_, &c. &c., may be explained, as already +remarked, by this process, rather than by fission, chorisis, or by +substitution of petals for stamens, &c. Each case must, however, be +examined on its own merits, as it is not safe to decide upon the +arrangement of parts in one flower by simply referring to the analogy of +others. In the following illustrations the course of development has +not, in all cases, been observed, and hence the explanation here given +must be taken with some reserve; for should it prove that the +adventitious lobes, &c., are formed simultaneously with the ordinary +petals, the case will be one of chorisis rather than of enation, as here +understood. Again, it may be that the supernumerary organs really +represent petals or stamens in disguise, though this hypothesis demands +the further assumption (in order to account for the interference with +the law of alternation) that suppression of certain organs has taken +place. + +Taking first those instances in which the supplementary petals appear on +the inner surface of the corolla, as being at once the most frequent, +and as presenting the closest analogy, with similar conformations, under +natural circumstances, certain double-flowered varieties of the Chinese +primrose, _Primula sinensis_, may be mentioned. In these flowers the +calyx is normal, the tube of the corolla is traversed by ten vascular +bundles, and the limb is divided into ten fimbriated lobes. About +halfway up the tube, on the inner surface, are given off five +supernumerary petals, opposite to as many lobes of the corolla. Some of +the supplementary petals have a stamen in front of them, in the same +relative position as in the normal flower. In some cases the back or +outer surface of the supplementary petal is turned towards the inner or +upper surface of the primary corolla, thus [Symbol: ((turned 90 degrees +cw]; while, in other instances, the front of the adventitious lobe is +directed towards the corresponding surface of the original petal, thus +[Symbol: () turned 90 degrees]. Whether these supernumerary petals are +formed by chorisis or by enation cannot, with certainty, be determined +without examining the early stages of development. + +[Illustration: FIG. 212.--_Datura fastuosa._ True corolla turned back to +show the supernumerary corolla with the petal-like segments attached to +its outer surface (reduced).] + +Of more interest are those instances where the adventitious growth is on +the outside of the corolla; thus in a garden azalea there was +intermediate between the calyx and the corolla, both of which were +normal, a series of five petalodes, alternating with the sepals, and, +therefore, opposite to the lobes of the corolla, and adherent with them +at the very base, though elsewhere detached. These petalodes were +concave on the surface looking towards the calyx, and were there +brightly coloured, while the tint of the opposite surface looking +towards the corolla was of a duller hue, corresponding with that of the +outside of the corolla-tube. This arrangement of the colour was thus +precisely similar to that which occurred in the four-winged leaves +already referred to at p. 446. In some flowers of _Datura fastuosa_ a +similar series of excrescences was observed; the calyx and the corolla +were normal within the latter, intervening between it and the stamens +was a second corolla produced by duplication, and adherent to the inner +surface of this latter were five stamens. So far there was nothing very +peculiar; it remains to say, however, that on the outer surface of the +second corolla were five petal-like lobes closely adherent to it below, +but partially detached above. The colour of the adventitious segments +was paler on the outside than on the inner surface, as in the corolla +itself. The position of the several parts was such that they were +opposite one to the other; hence, while the lobes of the inner corolla +were opposite to those of the outer one, the intermediate petalodes were +opposite to both; thus: + + S S S S S + -------------------------- + P P P P P + X X X X X + -------------------------- + P P P P P + st st st st st + +The X indicating the position of the petalodes. + +[Illustration: FIG. 213.--Gloxinia, with supernumerary segments on the +outside of the true corolla.] + +A still more singular case is that of a variety of the Gloxinia, +described originally by Professor Edouard Morren,[521] but which is now +becoming common in English gardens. When first observed these flowers +were observed to produce petaloid segments outside the ordinary corolla, +and partially adherent to (or rather, not completely separated from it) +much as in the azalea before mentioned, the outer surface being brightly +coloured, like the inner surface of the corolla in ordinary gloxinias. +Being encouraged and tended by gardeners, in course of time, instead of +a series of petalodes, more or less distinct from one another, a second +corolla or "catacorolla" was formed outside the primary one, so that a +hose in hose flower was produced, but, in this case, the supplementary +flower was formed on the outside and not within the ordinary corolla. +Moreover, the disposition of the colour was reversed, for in the +outermost corolla the richest hues were on the outer surface, while in +the inner or true corolla they were on the inside. + +Professor Morren considers the adventitious petalodes as rudiments of so +many supplementary flowers, axillary to the calyx, and adnate to the +corolla; each lobe then would, in this view, represent an imperfect +flower, and the completed catacorolla would be formed of a series of +confluent flowers of this description. But this view involves the +assumption of the suppression of all the parts of the flower, except the +lobes in question. + +[Illustration: FIG. 214.--"Catacorolla" of _Gloxinia_, formed from the +union of adventitious petalodes on the outside of the true corolla +(after Morren).] + +The view here propounded that the lobes in question are enations from +the true petals, which become confluent, so as to form the catacorolla, +is surely more simple, involves no assumptions of suppression of parts; +and moreover, is borne out by the examination of some flowers, where the +production of these adventitious lobes from the outside of the minute +partially developed petals could be distinctly seen. + +=Enation from the stamens.=--An illustration of this process occurred in +some double-flowered rhododendrons, which presented the following +arrangement of parts:--calyx and corolla normal; within the latter eight +petal-like stamens, forming a pseudo-corolla. The appearance presented +by the petaloid filaments and anthers was as if they were adnate to the +centre of the petals, but, on closer examination, it appeared that the +petaloid expansion to which the dilated filament was apparently +attached, was equally a part of the stamens; in other words, that the +filament was provided with four petal-like wings, two on each side +[Symbol: 00 topped by (turned cw 90 deg., an o above and another ( turned cw +above that]. This disposition was well seen in the anther, half of which +was, in some cases, petaloid like the filament; in fact, the inner wing +of the latter was directly continuous with the petal-like expansion from +the anther. A section through the latter showed, going from within +outwards, the cut edges of two perfect polliniferous lobes in the +centre; and on either side the petaloid wing representing the remaining +anther-lobe; outside these were the edges of the remaining wings, one on +each side. (See p. 290, fig. 155.) + +=Enation from the carpels.=--The only instances of this that need be +referred to are such cases as those in which spur-like projections, +horns, tubercles, or winged expansions, are formed from the surface of +the ovary during the course of its development. The extraordinary +cornute oranges described and figured by Ferrari, Gallesio, and other +writers on the genus _Citrus_, may be mentioned under this head. A +similar formation occurs in the fruit of some species of _Solanum_. (See +p. 316.) + +FOOTNOTES: + +[518] It is desirable in this place to allude to a singular case of +fissiparous division of a leaf of _Prunus Laurocerasus_ described by +Prof. Alexander Dickson ('Seemann's Journ. Botany,' vol. v, 1867, p. +323), and which did not come under the writer's notice till after the +sheet relating to fission, p. 61, had been sent to press. Dr. Dickson +thus speaks of this abnormal leaf:--"The petiole (unchanged) supported +two laminae, placed back to back, and united by their midribs (_i.e._ not +separated) to within about an inch from their extremities, which were +perfectly free from each other. These laminae stood vertically, their +edges being directed towards and away from the axis; and as they were +placed back to back, the shining surfaces, corresponding in structure to +the normal upper leaf-surface, were directed laterally outwards. In the +axil of this abnormal leaf were two axillary buds. The existence of two +leaf-apices and two axillary buds shows that this was not due to an +accidental exuberance of development, but to fissiparous division, +which, had it been complete, would have resulted in the replacement of a +single leaf by two leaves. The arrangement in Prof. Dickson's leaf may +be thus represented: [Symbol: )OO( with X above]. The nature of the case +may be even better seen by comparison with the normal arrangement, which +would be [Symbol: (OX turned 90 degrees ccw], while in those cases where +the fission of the leaf occurs in the same plane as that of the primary +lamina, as where a leaf splits into two lobes at the apex, with a midrib +to each, the arrangement is as follows: [Symbol: OX turned 90 degrees +ccw, with 2 arcs below forming a half circle], the X in all cases +representing the position of the axis, the O that of the axillary bud, +and the [Symbol: ( turned 90 degrees ccw] that of the laminae." + +[519] Linn., 'Phil. Bot.,' Sec. 274. The term "_crispa_" is surely +preferable to that of Re, "phyllorhyseme." + +[520] See C. Morren, "Consid. sur les deformations," &c., in 'Bull. +Acad. Belg.,' 1852, tom, xix, part 3, p. 444; and as to ferns, see +Moore, 'Nature-Printed British Ferns,' 8vo ed., where numerous +illustrations are given. + +[521] 'Bull. Acad. Belg.,' t. xix, p. 224, tab. i; and 'Gardeners' +Chronicle,' 1865, p. 865. + + + + +PART II. + +ATROPHY. + + +Under the head of atrophy are included those cases wherein the organs +affected are actually present, but in a dwarfed and stunted condition as +compared with surrounding parts. + +The diminished size is, in such instances, obviously due to a partial +development and to an arrest of growth at a certain stage, from the +operation of various causes, either external or inherent to the +organization itself. It may affect any part of the plant, and exists, in +very varying degree, in different instances, being sometimes so slight +in amount as not to preclude the exercise of the functions of the part; +while in others, the structure is so incomplete that the office cannot +be performed. These differences depend, of course, upon the stage of +development which the organ had reached when its growth was checked. For +practical purposes atrophy may be distinguished from suppression by the +fact that in the latter case a certain element of the flower or plant +which, under ordinary circumstances, is present, is entirely wanting, +while, in the former class, it exists but in a rudimentary condition. + +Again, atrophy is to be separated from that general diminution in the +size of the whole plant or of distinct parts of that plant which is +comprised under the term "nanism." Thus the several dwarf varieties of +plants (var. _nanae_), or those in which the leaves or flowers are +smaller than usual (var. _parvifoliae_, v. _parviflorae_), are truly +regarded as variations, and not as malformations properly so called. + +Atrophy is partial and special in its operation, nanism is general. + +Under ordinary circumstances atrophy is exemplified by the presence of +rudimentary or imperfect organs, as, for instance, in _Pentstemon_, +_Scrophularia_, &c., where one stamen is atrophied. + +For convenience sake atrophy may be divided into abortion and +degeneration, the first including cases where, from arrest of +development occurring at an early stage, organs are present; but in a +much smaller and more rudimentary condition than usual, their form and +general appearance, except so far as regards their dimensions, not being +materially altered. On the other hand, in cases of degeneration, +development is not entirely checked, but rather perverted, so that not +only the dimensions are lessened, but the form is altered. + + + + +CHAPTER I. + +ABORTION. + + +The sense in which this term is here understood has been explained in +the preceding paragraph. It is only necessary to say further, that cases +of abortion are to be distinguished from those of suppression, on the +one hand, and those of degeneration on the other. In suppression there +is from the first an absolute deficiency of a particular organ. In +degeneration the part is present, but in a diminished and perverted +condition. In abortion it exists, but in a stunted and dwarfed, but not +otherwise permuted state. + +=Abortion of axile organs.=--When the main stem is arrested in its +growth, the habit and general appearance of the plant are materially +altered, as in the so-called stemless plants, _plantae acaules_. In these +the internodes are so slightly developed that the leaves are closely +crowded in tufts or rosettes. When this shortening of the stem +(acaulosia) occurs, without other considerable change in other organs, +the deviation is classed under the head of variation rather than of +monstrosity; and, indeed, in very many plants, this arrested growth of +the axis is the rule rather than the exception. When occurring in an +abnormal manner, atrophy of the stem is most frequently attended by +other more or less grave alterations in other structures; thus +Moquin-Tandon[522] cites an instance of _Camphorosma monspeliaca_, +wherein the stems presented the form of very short, hard, woody +tubercles, thickly clothed with deformed leaves, and invested by a vast +number of hairs, longer and more dense than usual. A similar deformity +sometimes occurs in an Indian species of _Artabotrys_; in these +specimens the branchlets are contracted in length, and bear numerous +closely packed scaly leaves, densely hairy, and much smaller than +ordinary. + +Spines and thorns may he looked on as atrophied branches, and seem to +result from poorness of soil, as the same plants, which, in hungry land, +produce spines, develop their branches to the full extent when grown +under more favorable conditions.[523] + +In the birch an arrest of development in some of the branches is of +common occurrence. The branch suddenly ceases to grow in length; at the +same time it thickens at the end into a large bulbous knob, from which +are developed a profusion of small twigs, whose direction is sometimes +exactly the reverse of that of the main branch. (See p. 347.) + +The branches of the common spruce fir, especially the lateral ones, when +attacked by a particular species of aphis, are very apt to be developed +into a cone-like excrescence.[524] + +A shortened condition of the flower-stalks occurs occasionally, greatly +altering the general character of the inflorescence. This has been +observed in pelargoniums and in the Chinese primrose, in both of which +the effect was to replace the umbellate form of inflorescence by a +capitate one. + +=Abortion of the receptacle.=--Here may be mentioned those cases of +flowers with habitually inferior ovary (real or apparent), in which the +receptacle fails, from some cause or other, to dilate as usual. This has +already been alluded to under the head of Prolification, Displacements, +&c. (pp. 78, 130, &c., figs. 35-37, 64, &c.), and hence requires only +incidental comment in this place. There are, however, certain other +cases of a similar nature which may here be referred to; such as the +abortive condition of the inferior ovary, or rather of the receptacle, +that usually encircles the ovary in _Compositae_ and _Umbelliferae_. In +the former natural order the following plants have been met with in this +condition:--_*Tragopogon pratense!_, *_Cirsium arvense_, _Hypochaeris +radicata_, _Senecio vulgaris!_, _Coreopsis Drummondi_. In the latter +order, _Daucus Carota!_ _OEnanthe crocata!_ and _Thysselinum +palustre_, seem most frequently to have been observed in this +state.[525] In some gourds the receptacle may be seen partially +developed only, and forming a kind of cup, from which the true carpels +protrude. + +=Abortion of the leaves.=--Arrest of growth in the leaves occurs in +different ways; sometimes the whole leaf is smaller than usual; at other +times certain parts only are reduced in size; while, in a third class of +cases, portions of the leaf are entirely suppressed. + +Moquin[526] mentions having seen the leaves of _Chenopodium vulvaria_, +and of _Diplotaxis muralis_ reduced to a fourth of their natural size; +and he alludes to other cases of the same nature, seen by other +observers, in _Hypericum perforatum_ and _Blitum polymorphum_. + +_Nicandra physaloides_[527] has also been met with in a similar +condition, which, indeed, is a common result of insect-puncture, and of +fungous growth in plants. Those instances in which the leaf is +diminished in size, without any attendant malformation in other organs, +may be regarded rather as variations than as monstrosities, as in the +case of the entire-leaved varieties of those plants which ordinarily +have cut or divided leaves, _e.g._ _Plantago Coronopus_, var. +_integrifolia_, _Papaver Rhoeas integrifolia_, &c. &c. The same remark +may be made of those specimens in which one part of the leaf is +developed to a less extent than another, as happens in the submerged +leaves of such plants as _Ranunculus aquatilis_, _Cabomba aquatica_, the +spiney leaves of _Berberis_, the fenestrated leaves of _Ouvirandra_, &c. +In the illustrations last cited the relative deficiency of one portion, +as contrasted with another, takes place as a constant occurrence, and is +uniform and regular throughout the whole leaf. When, on the other hand, +the deficiency in question happens accidentally and irregularly, the +change may be considered as a malformation. One side of the blade of the +leaf is frequently affected in this manner, the other portions remaining +unaffected. It would appear as if any plant might be thus altered, but +the following species appear to be particularly subject to this change: +_AEesculus Hippocastanum_, _Digitalis purpurea_, _Morus alba_, _Fagus +silvatica contracta_ (hort.), _Codiaeum variegatum_ var. _erosum_ +(hort.), _Broussonettia papyrifera_, _Scolopendrium vulgare_, &c. + +Frequently this irregular diminution in proportion is coexistent with an +unusual degree of cleavage or laciniation of the margin, as in _Acer +platanoides laciniatum_, _Tilia asplenifolia_, _Alnus imperialis_ +(hort.), _Fagus silvatica_ var. (hort.), &c. + +In the case of what are sometimes termed interrupted leaves, the laminar +portions of the leaf are here and there deficient on both sides of the +midrib, leaving small portions of the latter, as it were, denuded and +connecting the segments of the laminae one with the other. This has been +observed amongst other plants in _Veronica latifolia_, _Broussonettia +papyrifer_, _Codiaeum variegatum_ var. _interruptum_ (hort.), +_Scolopendrium vulgare_, &c.[528] (See p. 328.) + +In some of the leaves which have been already referred to in +illustration of the inordinate growth of the cellular portions, the +increased development of parenchyma is associated with a contracted +state of the midrib and its branches, producing a puckered appearance of +the leaf, an exaggerated degree of that change which produces what are +termed "folia bullata." In illustration may be cited various species of +_Mentha_, _Perilla_, _Coleus_, _Fagus silvatica crispa_, _Cytisus_, +_Laburnum_ var., and other forms, cultivated in gardens for their +singularity. + +Entire absence of the stalk of the leaf occurs normally in sessile +leaves; on the other hand the blade of the leaf is only occasionally +developed in the phyllodineous Acacias, in some species of _Oxalis_, +_Indigofera_, _Lebeckia_, _Ranunculus_, _Bupleurum_, &c. + +De Candolle,[529] from a consideration of _Strelitzia juncea_, in which +the petiole alone is developed, was led to the inference that in many +monocotyledonous plants the blade of the leaf was never developed, the +portion present being the sheath or stalk, unprovided with limb. The +correctness of this inference is shown, amongst other things, by the +occasional presence of a leaf-blade in _Strelitzia juncea_ itself. + +Occasionally the laminar portions of the leaf are completely wanting, +leaving only the main ribs, as in the case of _Berberis_, while the +adjoining figure (fig. 215) represents an instance of a cabbage wherein +the innermost leaves are represented by thick fleshy cylindrical bodies +corresponding to the midribs of the ordinary leaves. There is in +cultivation a variety of the cabbage which constantly presents this +peculiarity. + +[Illustration: FIG. 215.--Inner leaves of cabbage reduced to their +midribs.] + +The suppression of one or more leaflets of a compound leaf has already +been referred to at p. 396. + +=Abortion of the perianth, calyx, and corolla.=--Illustrations of +partial development in these organs are not rare, under ordinary +circumstances, as for instance the "obsolete" calyx of Umbellifers. In +the cauliflower the branches of the inflorescence are contracted in +length, while their succulence is much increased; at their extremities +they bear crowds of imperfect flowers, in which the calyx only is +visible, and that only in a rudimentary and partially developed +condition. Imperfect development of the whole or of some of the +constituent parts is more common in the case of the corolla than in that +of the calyx. In _Arenaria serpyllifolia_ the petals, especially in +autumn, are only one fourth the length of the sepals. _Anagallis +phoenicea_, _Honckenya peploides_, _Arabis alpina_, _Ranunculus +auricomus_, _Rubus fruticosus_, and _Geranium columbinum_, also +frequently afford illustrations of this circumstance. + +[Illustration: FIG. 216.--Abortion of four out of five petals, _Viola +tricolor_, side and front views.] + +At fig. 216 is represented a pansy in which four of the five petals were +very small and colourless, while the lower spurred petal was of the +usual size and colour. In this flower the stamens and pistils were +wholly suppressed, and the flower-stalk, instead of being bent near the +flower, retained its primary straight direction. Similar atrophic +conditions of the corolla occur habitually among _Violaceae_. + +The diminished size of the petals sometimes coexists with an increase in +their number, as in a flower of _Streptocarpus Rexii_, mentioned by +Bureau.[530] + +Among monocotyledons this partial development seems to be even more +frequent than in dicotyledons. In addition to the well-known cases of +certain species of _Bellevalia_ and _Muscari_, wherein the uppermost +flowers of the raceme are more or less atrophied (see p. 347, fig. +179), a few less common illustrations may be cited. In crocuses it is +not a very uncommon circumstance to find the three inner segments of the +perianth smaller than natural, and generally unequal in size. This +occurs without any other perceptible change in the flower. + +Schlechtendal[531] mentions a flower of _Fritillaria imperialis_ in +which the perianthial leaves were relatively very small, and destitute +of the usual nectary, while the stamens, on the other hand, were of +their natural size and appearance. Fresenius[532] records a similar +occurrence in the same plant. + +Morren[533] gives details of like appearances in _Hymenocallis +americana_, and Delavaud[534] in _Tigridia pavonia_. + +In certain orchids an arrested development of the perianth is habitual, +as in _Oncidium abortivum_ (fig. 217), where, on a large branching +panicle, numerous abortive, but few perfect, flowers are produced. +In a similar way the petals and labellum of _Odontoglossum +Uro-Skinneri_ have been found reduced to filamentous processes. + +[Illustration: FIG. 217.--Flower of _Oncidium abortivum_, magnified.] + +=Abortion of the stamens.=--Atrophy of one or more stamens is of very +common occurrence, as a general rule, in many genera of plants, _e.g._ +_Scrophularia_, _Erodium_, many _Restiaceae_, &c. &c. As a strictly +teratological condition atrophy of the stamens is more rare than +complete suppression. It has been noticed in _Arabis alpina_, _Cerastium +glomeratum_, _C. tetrandrum_, _Rhamnus catharticus_, _Anemone_, +_Hepatica_, &c. It happens frequently among Orchids both wild and +cultivated. In the _Hymenocallis_ flowers described by the elder Morren, +four out of five stamens were atrophied. In other flowers, otherwise +perfectly formed, one abortive stamen was found bearing a spherical +indehiscent anther. All these atrophied anthers of _Hymenocallis_ were +found to contain pollen, differing at first sight but little from what +is usual, but presenting this important peculiarity, that while the +normal pollen does not burst until it comes into contact with the +stigma, in the abnormal flowers the outer coat of the pollen-grains +split while still within the anther, from which latter, indeed, they +could not escape, owing to the indehiscent nature of the latter. Again, +the pollen-tube of the abnormal grains cracked, in its turn, on mere +exposure to the air, and liberated the fovilla, so that the pollen of +these atrophied anthers was necessarily impotent, because it opened +before it could be applied to the stigma, even had that been rendered +possible by the opening of the anther. + +An abortive condition of the stamens and of the pollen, is of very +common occurrence among hybridised plants. Gaertner and other writers +have spoken of this defective condition as contabescence.[535] It forms +one reason for the sterility so frequently observed in the case of true +hybrids. In some hybrid passion-flowers, while all other parts of the +flower were apparently perfect, even to the ovules, the stamens were +atrophied, and distorted, and contained little or no pollen; the few +grains of the latter being smaller than usual. (See under Heterogamy, +pp. 193-196, and p. 398.) + +=Abortion of the pistil, fruit, &c.=--Traces of the carpels occur in +many male flowers of unisexual plants, _e.g._ _Sterculiaceae_, +_Euphorbiaceae_, _Restiaceae_, &c. &c., and in some natural orders there +appears to be a tendency towards a dioecious condition, _e.g._ +_Caryophylleae_, as in _Lychnis dioica_, _Silene otites_, _Arenaria +tetraquetra_, &c. The last-named plant is stated to have, in some cases, +imperfect pistils; in others, rudimentary stamens; while a third set of +flowers are hermaphrodite.[536] The ovary of aconites, according to +Moquin, is very subject to atrophy. + +[Illustration: FIG. 218.--Bladder plum.] + +During the maturation of the pistil, and its passage to the fruit, great +changes of consistence frequently take place, owing to the development +of cellular tissue, or of woody matter, according as the fruit is +succulent or woody. It sometimes happens that, owing to some disturbing +causes, the changes that usually occur fail to do so; thus, the stone of +plums is occasionally deficient, as in what are termed bladder-plums +(fig. 218); some of these, consisting merely of a thin bladder, are +curiously like the pods of _Colutea_.[537] + +MM. Fournier and Bonnet[538] describe a fruit of a _Rubus_, with +perfectly dry fruits, like those of a _Geum_, and this form was +considered by Steudel to form a distinct species. It is, however, merely +a variety in which the fruits have not become succulent.[539] + +Schlechtendal describes[540] the ordinarily baccate fruit of a vine as +becoming dry, and even dehiscing by valves like a capsule. + +In maize it occasionally happens that one or two of the longitudinal +series of fruits become abortive, leaving a smooth furrow, at first of a +greenish colour, but ultimately of a reddish yellow. Often a second row +of fruits, opposite to the first, is also atrophied, so that the whole +spike changes its cylindrical form for a flattened one.[541] See also +under Heterogamy, Meiophylly, &c. + +=Abortion of the ovules.=--In the case of a pluri-ovulate ovary it +rarely happens that all the ovules attain to maturity, some never get +fertilised, others, pressed on by their neighbours on either side, +become impeded in their development, and finally disappear, or remain as +rudiments.[542] This is the case, under ordinary circumstances, and +still more so in the case of hybrid plants, or of monsters. Where the +outer coats of the ovule become more or less leafy in appearance (see p. +262), the inner investments become more or less atrophied, or are even +more frequently entirely suppressed, as is also the nucleus. + +In other cases, a simple arrest of development takes place; the ovule, +for instance, which should be anatropal, remains straight, while the +integuments, checked in their development, form imperfect sheaths from +which the shrivelled nucleus protrudes. + +=Depauperated Ferns.=--The preceding illustrations have been taken from +flowering plants chiefly, but a similar defective development is +manifested in cryptogamous plants. The contraction and imperfect +development of the fronds of some varieties of ferns, hence called +depauperated, may receive passing notice, as also the cases in which the +sori or clusters of spore cases are denuded of their usual covering, +owing to the abortion or imperfect development of the indusium, as in +what are termed exindusiate varieties.[543] + +=General remarks on abortion, coincident changes, &c.=--Reference has +already been made, while treating of hypertrophy, suppression, &c., to +certain other changes affecting the flower at the same time. Atrophy of +one organ or set of organs, for instance, is frequently accompanied by a +compensating hypertrophy or by an increased number of other parts. In +the feather-hyacinth, _Muscari comosum_, var., _monstrosum_, the absence +of flowers is compensated for by the inordinate formation of brightly +coloured threads which appear to be modified pedicels (see pp. 347, +348); so also in the wig plant, _Rhus Cotinus_. So the atrophy of the +stamens, in some flowers, is coincident with the hypertrophy of the +pistil. Thus, Unger, 'Denkschr. d. Kais. Acad. der Wissensch. Math. Nat. +Classe,' Mai 25, 1848, p. 103, tab. ix, describes a case wherein the +corolla and stamens of _Desmodium marylandicum_ were atrophied, while +the calyx and legume, on the other hand, were hypertrophied. + +Fusion of the members of one whorl with one another, or with the +components of an adjacent series, often entails atrophy or suppression, +either in the united organs themselves, or in adjacent ones. A +foliaceous condition of the outer portions of a flower is very generally +attended by atrophy or complete suppression of the inner portions. + +From this point of view the observations of Morren[544] on the different +degrees of atrophy up to complete suppression, observable in the flowers +of _Bellevalia comosa_, are of importance. According to this observer, +the most highly differentiated parts, such as the stigmas, the ovules, +and the anthers, are the first to disappear, the filaments often being +developed without anthers. Ultimately a deformed and empty perianth +alone remains. In the ordinary course of things the mouth of the +perianth is open, but in some of these malformations it is closed, and +when that happens, the effects of atrophy are the more observable in the +stamens and pistils. + +The impotence of the pollen in certain atrophied flowers, as noticed by +the same observer, is of much interest, especially in reference to the +sexual relationship between the different forms in polymorphic flowers +as studied by Mr. Darwin. + +A change in direction may also be noted as a common accompaniment of +atrophy or suppression; thus, in a capsule of _Veronica Beccabunga_, +which was one-celled by the abortion of one carpel, the style was +lateral instead of terminal. + +As to the causes of these structural deviations but little is known; +certain of them have been already alluded to. In some cases atrophy and +suppression maybe regarded as permanent states of a condition usually +transitory, but this is clearly not always the case. Among external +causes anything bringing about an enfeebled condition might be supposed +to lead to atrophy, or suppression of some parts. + +Gaertner[545] attributes the arrested development and fall of flowers to +some among the following causes:--1. non-application of the pollen of +the same variety, and consequent imperfect fertilisation; 2. any +considerable injury to the calyx, &c.; 3. destruction of the style or +stigma before the fertilisation of the ovary; 4. application to the +stigma of imperfect or heterogeneous pollen or indifferent pulverulent +matter; 5. defective conceptive power in the ovary. + +Abortion of the ovules is considered by the same authority to be due +to--1. deficiency of heat; 2. excess of moisture; 3. peculiar formation +of the ovary; 4. over-luxuriant development of roots or buds; 5. +peculiar conditions of cultivation; thus, cuttings and layers produce +sterile and abortive seeds much more frequently than plants of the same +species raised from seed; 6. abortion of the seed is often combined with +luxuriant development of the walls of the fruit. + +Temperature and climatal changes in general seem not to be without +effect, as has been already mentioned in the case of _Arenaria +tetraquetra_, which is polygamous when growing in mountain districts. +Other illustrations of a similar character are mentioned under the head +of Heterogamy (p. 196). + +Pressure has been already alluded to as one of the most obvious of the +inducing causes of atrophy and suppression. + +In the case of _Ranunculus auricomus_ before cited, in which the petals +are rarely perfect, M. de Rochebrune considers that the deficiencies in +question depend, in great measure, on the amount of moisture in the +localities where the plant grows. In most places the flowers and carpels +are apt to become more or less abortive, while the leaves are luxuriant; +while, in dry places, the foliage is small, but the flowers are more +perfect. This is quite consonant with other facts relating to the +development of flowers or of leaves in general. + +But while external agencies undoubtedly play some part in bringing about +these changes, it is almost certain that internal causes inherent to the +organization of the plant are more important. Mr. Darwin[546] accounts +for the existence of rudimentary organs by the operation of the general +rule of inheritance, and explains their stunted condition as the effect +of disuse, not so much, of course, in the particular flower as in its +predecessors. This disuse may be the result of the superior efficacy of +foreign pollen as contrasted with that formed in the individual flower +itself. In this way many hermaphrodite flowers tend to become +dioecious, as in _Caryophyllaceae_, _Orchidaceae_, _Plantaginaceae_, +_Primulaceae_ and other orders. + +Although many of the circumstances above mentioned apply to plants whose +structure is habitually rudimentary, there is no reason why they may +not, under due restrictions, be applied to plants whose organs are only +occasionally defective. + + For further remarks on the subject of Abortion, the reader is + referred to the sections relating to suppression, etc., also to + Moquin-Tandon, 'El. Terat. Veget.,' p. 120; C. Morren, "De + l'atrophie en general," in 'Bull. Acad. Belg.,' t. xviii, 1851, + part i, p. 275. + +FOOTNOTES: + +[522] 'El. Ter. Veg.,' p. 132. + +[523] _Spinosae arbores cultura saepius deponunt spinas in hortis_, 'Linn. +Phil. Bot.,' Sec. 272. + +[524] Mr. Selby, in his 'History of British Forest Trees,' p. 465, gives +the following account of the formation of this peculiar growth:--"In the +autumn the parent aphis deposits her eggs at the base of the embryo +leaves, within the bud destined to produce the shoots of the following +year. When these begin to burst and expand in spring, the leaves, at +whose bases the eggs have been deposited, instead of increasing in +length, enlarge at the base, and form a cell or cyst whose mouth is at +first closed by a red velvety-looking substance. If opened in this state +a nest of small greenish aphides is distinctly visible, and at a certain +period, or when they have acquired maturity, which is towards the end of +the summer, the mouth of the cell opens and the insects fly off to +inflict a similar injury upon the nascent buds of the year. In some +instances the leaves of only a portion of the circumference of a shoot +are affected, in which case, though a slight distortion may take place, +the branch is not prevented from elongating; but in others, where the +whole of the leaves around the shoot are converted into nidi, elongation +is prevented and distortion to a great extent takes place." + +[525] See Cramer, 'Bildungsabweich.,' pp. 53, 64, for further +references. + +[526] 'El. Ter. Veg.,' p. 124. + +[527] Schlechtendal, 'Bot. Zeit.,' 1857, vol. xv, p. 67. + +[528] On the subject of this paragraph the reader may consult A. Braun, +"Ueber abnorme Blattbildung," &c., in 'Verhandl.,' d. 35, +'Naturforscherversammlung;' Jaeger, 'Flora,' 1850. p. 481, tab. 4, +_Digitalis_. + +[529] 'Org. Veget.,' i. p. 286. + +[530] 'Bull. Soc. Bot. France, vol. viii, 1861, p. 710. + +[531] 'Linnaea,' 1830, vol. v, p. 492. + +[532] 'Mus. Senkenb.,' ii. p. 45. + +[533] 'Bull. Acad. Roy. Belg.,' 1851, t. xviii. part i, p. 275. + +[534] 'Bull. Soc. Bot. France,' vol. viii, 1861, p. 147. + +[535] See Darwin, 'Variation of Domest. Anim. and Plants,' ii, 165. + +[536] Gay, 'Ann. Sc. Nat.,' ser. i, 1824, t. iii, p. 44. + +[537] See De Candolle, 'Mem. Legum.,' tab. 3, f. 1; Wyville Thomson, +'Trans. Bot. Soc. Edinb.,' 1851, July 10th; Berkeley, 'Gardeners' +Chronicle,' June 22nd, 1867, p. 654. A similar case is described by Dr. +Robb, in Sir W. Hooker's 'Journal of Botany,' 1841, vol. iii, p. 99, +with illustrative figures. The specimens there described were produced +at New Brunswick, where plum trees flower very freely, but seldom +produce ripe fruit. Dr. Robb's account is as follows:--"In the summer of +1839 I had an opportunity of watching the process of destruction among +the plums, and it was as follows--Before or soon after the segments of +the corolla had fallen off, the ovarium had become greenish yellow, +soft, and flabby. As the fruit continued to increase in magnitude, its +colour grew darker and of a more ruddy yellow, and at the end of a +fortnight or three weeks the size of the abortive fruit rather exceeded +that of a ripe walnut. In fact, an observer might imagine himself to be +walking amongst trees laden with ripe apricots, but, like the fabled +fruit on the banks of the Dead Sea, these plums, though tempting to the +eye, when examined, were found to be hollow, containing air, and +consisting only of a distended skin, insipid, and tasteless. By-and-bye +a greenish mould is developed on the surface of the blighted fruit; then +the surface becomes black and shrivelled, and at the expiration of a +month from the time of flowering the whole are rotten and decomposed. +The flower appears about the beginning of June, and before August there +is hardly a plum to be seen. It is curious that where two flower-stalks +arise from one point of the branch, one will often go on to ripen in the +normal way, while the other will become abortive, as above described." + +In a specimen described by Mr. Berkeley there were two distinct ovules +of equal size close to the apex of the fruit, connected with the base by +vessels running down the walls. It should be observed that there is a +worthless variety of plum, Kirke's stoneless, or Sans Noyau, in which +the kernel is not surrounded by any bony deposit. + +[538] 'Bull. Soc. Bot. Fr.,' 1862, vol. ix, pp. 37 et 291. + +[539] Carl Schimp, 'Fl. Friburg,' vii, p. 745; Hook, fil., 'Journ. Linn. +Soc.,' vi, p. 9. + +[540] 'Linnaea,' vol. v, 1830, p. 493. + +[541] Moquin-Tandon, 'El. Ter. Veg.,' p. 325. + +[542] Alph. De Candolle states that the position of the abortive ovules +affords a good character for discriminating between certain species of +_Quercus_, 'Bibl. Univ. Genev.,' 1862, t. xv, p. 929. + +[543] See Moore, 'Nature-Printed Ferns,' 8vo, for numerous illustrations +both of depauperate and exindusiate ferns. _Scolopendrium vulgare_ seems +to be one of the ferns most commonly affected in this way. Moore, loc. +cit., vol. ii, pp. 135, 147, 159, 165, &c. + +[544] 'Bull. Acad. Belg.,' t. xvii, p. 38, t. 1; Lobelia, p. 85. + +[545] Cited in 'Henfrey's Botanical Gazette,' i, p. 179. + +[546] 'Origin of Species,' p. 450. + + + + +CHAPTER II. + +DEGENERATION. + + +While the terms atrophy and abortion apply in the main to a mere +diminution of size, as contrasted with the ordinary standard, +degeneration may be understood to apply to those cases in which not only +is the absolute bulk diminished, but the whole form is altered and +depauperated. Degeneration, thus, is the result not so much of a +deficiency in growth as of a perversion of development. + +Under natural, _i.e._ habitual circumstances, the formation of pappus +in place of a leafy calyx may be considered as an illustration of +degeneration. It is evident, however, that no very decided line of +demarcation can be drawn between cases of perversion and of arrest of +development. + +=Formation of scales.=--These may be mere epidermal excrescences, or +they may be the abortive rudiments of leaves. Of this latter nature are +the "cataphyllary" leaves which invest the root stocks of so many +perennial plants, the perulae of leaf-buds, or the paleae on the common +receptacle of composite flowers. Other illustrations of a like character +are to be met with in the membranous scales that represent leaves in +_Ruscus_, _Asparagus_, _Pinus_, &c. Similar productions are met with +within the flower, where they may occur as the representatives of +sepals, petals, stamens, or pistils, or as mere excrescences. (See +Enation.) Whole families of plants, _e.g._ _Sapindaceae_, are +characterised by the presence of these organs, which are often of great +interest to the morphologist as indicating the true symmetry of the +flower, while they have acquired fresh importance since the publication +of Mr. Darwin's work on the 'Origin of Species,' wherein we are taught +to regard these rudiments as, in many cases, vestiges of organs that +were more completely developed in the progenitors of the present race of +plants, and the exercise of whose functions, from some cause or other, +having been rendered impossible, the structures become, in process of +time, proportionately stunted. + +Thus, in dioecious plants we frequently find traces of stamens in the +female flowers, and rudiments of the pistil in the male flower, +indicating, according to the Darwinian hypothesis, that the ancestors of +these plants were hermaphrodite (see Heterogamy). + +Mr. Darwin has also shown that, in some cases, the utmost degree of +fertility is attained, not from the action of the pollen on the stigma +of the same flower, but on the influence of the male element of one +blossom upon the female organs of another flower on another individual +plant. + +Hence, in such plants there is a tendency to a separation of the sexes, +while, from what has been before stated, it might be expected that +rudiments of the male or female organs would be found, and also as a +result of the operation of the law of inheritance. On the same +principles it is easy to understand the occasional presence of the +perfect in place of the rudimentary organs, as in _Dianthus_. + +In some instances the assumption of a scale-like form by any organ is +attended by a change in texture, the organs becoming dry and scarious, +or fleshy. Moquin cites in illustration of the first phenomenon the +flower of a _Vicia_, in which the petals were thick and fleshy, like the +scales of a bulb; and of the second the leaves of a _Chrysanthemum_, +which were replaced by small, glossy scales, like those which invest +ordinary leaf-buds. Sometimes the entire flower is replaced by +accumulations of small, acute, green scales. Cases of this kind, wherein +the flowers of a pea and of the foxglove were replaced by collections of +small ovate green scales packed one over the other till they resembled +the strobile of a hop, have been already alluded to. Most of these +scales are represented as having had other accumulations of scales in +their axils. + +Similar collections of scales may frequently be met with in the birch +and in the oak, and probably represent abortive leaf-buds. Other cases +of a like kind in _Gentiana Amarella_, where the scales are coloured, +are mentioned elsewhere. + +In some kinds of _Campanula_ a similar change is not uncommon. + +=Formation of hairs, spines, &c.=--The adventitious production of hairs +is likewise frequently due to an arrested growth, in some cases arising +from pressure impeding the proper development of the organ. In other +cases the formation of hair seems to accompany the diminished +development of some organ, as on the barren pedicels of the wig plant, +_Rhus Cotinus_. A similar production of hair may be noticed in many +cases where the development of a branch or of a flower is arrested, and +this occurs with especial frequency where the arrest in growth is due to +the puncture of an insect, or to the formation of a gall. In such cases +the hairs are mere excrescences from the epidermis. + +Prickles differ but little from hairs save in their more woody texture, +but true spines or thorns are modifications either of a leaf or of a +branch. Their presence seems often dependent on the soil in which the +plants grow, or on other external circumstances. + +They occur normally in the sepals of _Paronychia serpyllifolia_ and +other plants. + +=Formation of glands.=--Under this name are associated a number of +(generally) rudimentary organs very different in their morphological +nature and significance, and also in their functions. Some are truly +glandular or secreting organs, while others have no visible office. +Anything like a complete account of these structures would be out of +place, and reference is only made to them here on account of the +occasional existence of intermediate forms, which throw light on the +morphological significance of these structures. Thus, in _Passiflora_ +and _Viburnum Opulus_, the so-called glands on the sides of the petiole +appear to represent leaflets, and are not unfrequently developed as +such. + +M. Dunal observed a flower of _Cistus vaginatus_ in which some of the +stamens were replaced by an hypogynous disc.[547] Moquin has seen +similar instances in the flowers of a Rose, _Hypericum_, and Poppy. + +M. Planchon[548] gives an account of some very curious malformations in +_Drosera intermedia_, which go to show that the ovules are homologous +with the glandular hairs on the margins of the leaves of these plants, +an opinion corroborated by the researches of MM. Groenland and +Trecul.[549] + +Dr. Hooker shows that the pitcher of _Nepenthes_ is due to a +modification of a gland placed at the extremity of the midrib.[550] + +=Formation of tendrils.=--These are of very varied morphological import; +sometimes they are degenerated peduncles, as in passion-flowers, or +vines; at other times they are of foliar origin; or, again, they may +proceed from the segments of the perianth, as in _Hodgsonia_ and some +other cucurbitaceous plants. From their very different origin in +different plants it is necessary to study the development in each case, +and not apply to the generality what may be peculiar to one. In any case +this formation in question generally belongs more to general morphology +than to teratology.[551] + +Kirschleger, however, has recorded the existence of a cirrhose sepal in +_Cucurbita Pepo_.[552] + +FOOTNOTES: + +[547] 'Consid. Org. Fleur.,' p. 44, pl. ii, fig. 23. + +[548] 'Ann. Sc. Nat.,' 3 ser., Bot. ix, pl. 6, ff. 1, 2. + +[549] 'Ann. Sc. Nat.,' 3 ser., Bot. 1855, pp. 297, et 303. + +[550] 'Trans. Linn. Soc.,' xxii, p. 415. + +[551] See Darwin, "On Climbing Plants," 'Journal of Linnean Society,' +vol. ix, p. 1. + +[552] 'Flora,' 1845, p. 615. + + + + +GENERAL CONCLUSIONS. + + +At the end of many of the preceding sections, and whenever the +requirements of the case demanded it, a brief summary of the main facts +and of the inferences to be derived from them has been given. It may be +useful to give in conclusion a few general remarks on the whole subject. + +It will be seen from the numerous facts herein cited, that the so-called +monstrous formations (excluding morbid growths the result of disease or +injury) present no peculiarities absolutely foreign to the normal +organisation of plants. The difference between the natural and monstrous +development is one of degree and frequency of occurrence, not of kind. + +Deviations from the customary form have been shown to arise from +excessive or diminished growth, or from arrested or exalted development. +Even in those instances where, for convenience' sake, the term perverted +development has been used, it must be understood as applying only to the +particular plant or organ under consideration, as the form assumed is +perfectly in accordance with the ordinary conformation of some other +plant or group of plants. + +The period at which malformations occur is a matter of some importance; +this is, indeed, implied in the term arrest of development; evolution +goes on with growth up to a certain point and is then stopped, and thus +changes are brought about in the part affected of a different nature +from those dependent on non-development or suppression. + +Some malformations are congenital, therefore, while others are +acquired--in the former instance the disturbance is coeval in origin, +and contemporaneous in its growth and development, with those of the +affected part; in the latter case the organ may have attained its +ordinary degree of perfection, or at least may have advanced some way +towards it, before any deviation shows itself. True chorisis or fission, +for instance, is usually a congenital affection, arising at a very early +period of development, while enation takes place from structures which +are all but complete as to their organisation, even though they may not +have attained their full dimensions. The date of appearance is also of +consequence in determining the true nature of some changes; it does not +always follow, for instance, that because one organ occupies the +position of another, it is of the same nature as the one whose place it +fills. The presence of anthers on petals or on such organs as the corona +of _Narcissus_ does not necessarily constitute those parts actual +stamens, but rather staminodes. The true stamens are either wanting, or +if present, they are in advance of their imitators as regards their +development. + +=General morphology of the leaf and axis. Homology.= Since the time when +Goethe's generalisations were adopted by A. P. De Caudolle, special +attention has been given to the form and mode of development of the +leaf-organ; for as it was well said by Wolff, if once the course of +evolution and the structure of the leaf were known, those of the parts +of the flower would follow as a matter of course. + +It is not necessary, in this place, to pursue the subject of the +development and construction of the leaf further than they are +illustrated by ordinary teratological phenomena. + +From this point of view perhaps the most interesting circumstance is the +part that the sheath of the leaf plays.[553] In many cases of so-called +metamorphosis, it is the sheath of the leaf that is represented and not +the blade. In normal anatomy the sepals, petals, carpels, and even the +stamens, as a general rule, correspond to the sheath rather than to the +blade of the leaf, as may be seen by the arrangement of the veins. The +blade of the leaf seems to be set apart for special respiratory and +absorbent offices, while the sheath is in structure, if not in office, +more akin to the stem. It would not be easy apart from their position to +distinguish between a tubular sheathing leaf and a hollow stem. The +development of adventitious growths by chorisis or enation has been +frequently alluded to in the foregoing pages, and many illustrations +have been given of the power that leaves have of branching in more than +one plane, owing to the projection of secondary growing-points from the +primary organ. These new centres of development are closely connected +with the fibro-vascular system of the leaf, so that no sooner does a new +growing point originate, than vessels are formed to connect the new +growth with the general fibrous cord, see pp. 355, 445. This leads M. +Casimir De Candollo to consider the entire leaf as a composite +structure. The morphological unit, says he, is the cellular protrusion +or growing point (_saillie_) and its corresponding fibro-vascular +bundle.[554] + +The identity, in a morphological point of view, of the leaves and the +lateral parts of the flower is so thoroughly recognised that little +need be said on that score, save to repeat that the homology of the +floral organs is usually not so much with the entire leaf as with its +sheath. + +The most singular instances of morphological identity are those relating +to the sexual organs. We have seen the gradual transition of stamens to +pistils, and of pistils to stamens, the development of ovules on the +edges of the anther, the co-existence of pollen with ovules on an +antheroid body, and, stranger still, the actual development of pollen +within the tissues of the ovule itself! From such facts, in addition to +what we know of the relative position, internal structure, and mode of +development of the organs, it is impossible to avoid coming to the +conclusion that, however distinctly these parts may, under ordinary +circumstances, be set apart for the performance of distinct functions, +morphologically they are homologous. + +These ideas may be carried yet farther--the same sort of evidence, which +is adduced in support of the morphological identity of leaves with the +parts of the flower, may be advanced in confirmation of the opinion, +that, morphologically, there is no distinction between axis and leaf. +The leaf, according to this view, is a specialised portion of the axis +set apart to do certain work, just as the petals, stamens, &c., are +leaves told off for distinct uses. It is unnecessary to refer to the +intermediate productions linking the leaf-form to that of the axis, all +that is requisite here is to point out the facts that teratology lends +in support of these views. These may be summed up by the statement that +almost all those attributes which morphologists recognise as peculiar to +one or the other organ respectively, may be and are manifested by both. +We have the stem acquiring the characters of the leaf, and the leaf +those of the stem. Thus we have seen leaves, leaf-buds, branches, and +flower-buds springing from leaves or leaf-organs;[555] see pp. 174, 177, +445, &c. The structure that we are apt to associate exclusively with +one is found to pertain to the other. The arrangement of the vascular +cords in the leaf-organ finds its counterpart in the axis, generally, it +is true, modified to suit altered circumstances or diverse purposes. In +some cases the disposition is absolutely indistinguishable in the two +organs. It may then be said that the distinctions usually drawn between +axis and leaf are not absolute, and that, however necessary such a +separation may be for descriptive or physiological purposes, +morphologically the two organs are identical. Again, it may be said that +leaf and axis are two phases of the same organ,--an organ capable of +existing in its undifferentiated state in the form of a thallus among +Cryptogams, but which in the higher groups of plants becomes marked out +into separate portions, each portion having its own distinct functions +to fulfil for the common benefit of the whole organisation.[556] + +=Special morphology.=--Under this heading brief reference may be made to +some of the organs whose morphological nature has been, and still is, +much contested. It is clear that for the due elucidation of these +matters, development and the comparative investigation of similar +structures in different plants must be studied. Teratological data by +themselves can no more be trusted to give a correct solution of any +particular question, than the evidence furnished by other departments of +botanical science taken separately. With this statement by way of +caution, allusion may be made to some of the organs whose morphological +construction is illustrated by the facts recorded in the present +volume. + +=Calyx-tube.=--In descriptive botany it is the common practice to speak +of a calyx-tube, by which is meant a tubular or sheathing portion at the +base of the flower, below the sepals or calyx-lobes, and distinct or +inseparable from the ovary. The question morphology has to solve is +whether this tubular structure is to be considered as a portion of the +axis, or whether it is to be regarded as composed of the confluent bases +of the sepals. + +Mr. Bentham, who has recently reviewed the evidence as to the nature of +the calyx-tube in his paper on _Myrtaceae_,[557] still holds to the +notion that the "calyx-tube" or "hypanthium" is formed from the +concretion of the basal portions of the sepals. He founds his +conclusions upon such facts as the following: the circumstance that the +point of origin of the leaf is not always the same as the point of +disarticulation or separation from the axis, inasmuch as the basal +portion of the leaf is often adherent to the stem for some distance, +though still recognisable as foliar not axial in its nature. In the same +manner, the corolla and androecium may be concrete at the base, so +that the stamens are for convenience' sake described as inserted into +the tube of the corolla, though it is generally admitted that both +stamens and petals are really hypogynous, and it is not usual to +consider the corolla-tube up to the divergence of the stamens as part of +the receptacle. A similar remark applies to the carpels and placentas. +Mr. Bentham further considers that the gradual disconnection of the +various whorls, that may be traced in many plants, is a further proof of +concretion, rather than of expansion of the axis, but this argument may +fairly be met by the consideration that the several whorls emerge at +different heights.[558] + +Organs originally free and distinct become ultimately combined at the +base by the gradual protrusion from the receptacle of a ring or tube +under them, as in the stamens of _Leguminosae_; yet, says Mr. Bentham, no +one would propose to describe the staminal tube of monadelphous +_Leguminosae_ as part of the receptacle and not of the stamens. Perhaps +not, for descriptive purposes, but morphologically it would not be easy +to separate such a tube from the receptacle. The principal kinds of +malformation which have a bearing on this subject are mentioned at pp. +77-81 and 247, from which it may be seen that the evidence furnished by +teratology is conflicting. It would seem, indeed, that while in some +families of plants there may be a real calyx-tube, in others the tubular +portion is a sheath-like prolongation of the axis. In _Primula_ or +_Pedicularis_, where the venation is clearly laminar, the tubular +portion is distinctly calycine. In other cases the so-called calyx-tube +seems as certainly to be an expansion of the receptacle, as in +_Rosaceae_, _Myrtaceae_, _Melastomaceae_, _Passiflora_,[559] &c. + +Where the petals and stamens are described as being inserted into the +throat of the calyx, or are perigynous, it may be assumed as a general +rule, subject to but few exceptions, that the so-called calyx-tube is +really a portion of the receptacle.[560] After all, this is very much a +question of words, and for the following reasons,--very often the base +of the calyx does evidently form a tube, and no one can say where the +calyx ends and the receptacle begins. Again, many leaves are known to +originate in the form of a ring-like protrusion from the axis, and from +this primary ring originate secondary developments. Thus the asserted +difference between a leaf, with such a history of development, and an +axial structure becomes obliterated. From this point of view, peltate +leaves like those of _Tropaeolum_ or _Nelumbium_ become very significant. +In both the leaf-stalk is cylindrical and traversed, as in the case of +all cylindrical leaf-stalks, by a circle of fibro-vascular cords, as in +a branch, and which radiate in all directions in the blade of the leaf. +Now, if (as often happens to a slight extent) the central portion of the +leaf were much depressed, owing to the disproportionate growth of the +peripheral, as contrasted with the central portions, we should have a +funnel-like or tubular formation, precisely similar to many of the +so-called calyx-tubes. And, if we further suppose new growths to +originate from the sides of this funnel or tube, by chorisis or enation, +we should have the homologue of a tubular calyx, to the inner surface of +which are attached petals, stamens, &c. From the consideration of +circumstances such as these just detailed, together with that of the +arrangement of the vascular cords, M. Casimir De Candolle arrives at the +conclusion that the calyx-tube is a ring-like projection from an axis +whose further direct development is arrested. The secondary projections +or growing-points correspond to the several fibro-vascular cords of the +primary ring, and are ultimately developed into sepals, petals, stamens +and ovaries (see pp. 394, 509). + +=Androecium.=--The main points of morphological interest relating to +the androecium, referred to in this volume, are those concerning the +structure of the anther (see p. 292), the compound nature of the stamens +in some orders (see pp. 294, 345), and the nature of the androecium in +orchids (see p. 380). + +=Inferior ovary.=--Is the pistil always foliar in its morphological +nature, or is it, in some cases, as Schleiden taught, formed from the +axis alone? To a great extent the reply to this question is dependent on +the conclusions that may be arrived at as to the true nature of the +calyx-tube. Considered from a teratological point of view, there is no +reason for considering the inferior ovary to be purely axial. On the +contrary, the evidence derived from this source supports the ordinary +opinion that the carpels are invaginated within the expanded top of the +flower-stalk and more or less adherent to it. Some of the gourds afford +good illustrations of this, the upper part of the carpels in these +fruits projecting beyond the axial portion. But this matter loses much +of its importance if the morphological identity of axis and leaf-organ +be conceded. The carpels in inferior ovaries seldom or never correspond +to the lamina of the leaf, and between the vaginal portion of the +carpellary leaf, and the axis who shall draw the distinction? + +=Placentation.=--Some botanists have considered the placentas to be +portions of the carpel, and have compared the production of ovules on +them to the formation of buds on the leaf of _Bryophyllum_. Others have +been led to see in each placenta, even when it is, to all outward +appearance, a portion of the carpellary leaf, a direct prolongation from +the axis, adherent to the leaf. Teratology shows that ovules may be +formed indifferently on leaf-organs or on stem-organs. Sutural, +parietal, axile, free-central placentation, and, if there be more forms, +all may be met with even in the same ovary (see pp. 96, 508). Now, if +there were such special tendencies in the axis, as contrasted with the +leaf, to produce ovules, it is hardly likely that such anomalous +arrangements as those just mentioned would be as frequent as they are. +But as leaves produce other leaves, from their edges or their surfaces, +and as they form buds in the same situations, just as axial organs +do,[561] there is surely little ground for considering the placentas, or +ovuliferous portions of the plant, to be of necessity axial. Here again, +much of the difficulty vanishes if the morphological identity of the +leaf-form and of the stem-form be admitted. + +=Structure of the ovule.=--The nature of the ovule and of its coverings +has been a fertile source of controversy. The teratological data bearing +on this subject have been given at pp. 262-272. These data strongly +support the notion of the foliar nature of the coatings, and of the +axial nature of the nucleus, taking leaf and axis either in the ordinary +sense, or as modifications one of the other. It has been shown that the +ovular coats may themselves become carpels, and that ovules may be +developed upon ovules, p. 268. Whether the intra-carpellary siliques of +_Cheiranthus_, not uncommonly met with (p. 182), are instances of ovular +transmutation may be open to doubt. + +The axial nature of the nucleus has been inferred from its position, +mode of growth, and from its occasionally lengthening into a leafy or +even a floriferous shoot. Probably it may occasionally be invested by +sheathing coats, more analogous to tubular processes from the +receptacle, than to foliar organs, as is the case in _Welwitschia_. The +discussion of this matter, however, pertains rather to normal morphology +than to teratology. + +=Morphology of conifers.=--The nature of the pseudo-leaves of +_Sciadopitys_, and probably of other Conifers, is illustrated by +teratology, as also is the true constitution of the scale of the cone +(see pp. 192, 245, 352), though it must be admitted that little or no +light is thrown on that much-contested point--the true nature of the +ovule of Gymnosperms. + +=Relative position of organs.=--When organs are considered, not +separately, but in their relations to each other, the appearances +presented are referable to similar causes. Thus, the separation of parts +usually united has been shown to depend on an excess of development, +the persistent union of parts, usually separate in the adult state, has +been traced to an arrest of the process of development, by no means +necessarily coexistent with diminished growth. The diminished or +increased number of parts is, in like manner, attributable to analogous +causes, as also are the variations in arrangement and form, spoken of +under the heads of Displacement, Peloria, Substitution, &c. + +In the instance of displacements, it has been shown how slight a change +is required to transform the so-called inferior ovary into a superior +one. A defective development of the top of the flower-stalk in some +cases, in others a lack of union between the tube of the receptacle or +of the calyx (comprising in those terms not only the apex of the +receptacle, but the base of the sepals) and the carpels, suffice to +bring about this change in a character which for systematic purposes is +of great value. + +=Law of alternation.=--The circumstances that interfere with the law of +alternation may be briefly alluded to. The deviations from the customary +arrangement have been very generally attributed to suppression, or to +chorisis. It is unquestionable that either of these affords an efficient +explanation of the arrangement in question, as also does that +modification of chorisis, as it may be considered, which has been +treated of under the head of Enation. Spiral torsion of the axis would +likewise bring about analogous results. Still, it is quite conceivable +that opposition or superposition of organs may occur without the +intervention of any such operations. This will be the more readily +conceded when it is remembered that the phyllotaxis of leaves not +unfrequently varies on different branches of the same individual tree, +and that a similar variation in the flower would at once disturb the +customary alternate arrangement. Coalescence of the vascular bundles in +an unusual manner, and an irregular disposition of these cords have +also been considered to bring about deviations from the rule of +alternation, but in general the formation of the cords is subsequent to +that of the growing points or mamelons. + +Adhesions, accompanied by displacements, occasionally produce similar +deviations, the nature of which is usually easily detected. + +=Co-relation.=--The importance of this subject first prominently brought +into notice by Geoffroy St. Hilaire gains in force daily. Rarely is a +malformation an isolated phenomenon, almost always it is associated, +from the operations of cause or effect, with some others. Instances of +this co-relation have been cited in the preceding pages, and many more +might have been mentioned, had the consideration of the relationship +between form and function formed part of the plan of this volume. A +change in itself slight, often acquires importance from its association +with other alterations. This is particularly well seen in the case of +the receptacle. Let an ordinarily concave thalamus remain, from +defective development, flat, and how great the change in the appearance +of the flower. Let the usually contracted receptacle be lengthened, and +the whole aspect of the flowers so affected is altered to such an extent +that, were their history not known, botanists would have no hesitation +in assigning them to widely separate groups in their schemes of +classification. Peloria, too, of either form, affords excellent +illustrations of the co-existence of one changed condition with another. +Not only is the form of one set of organs altered, but the number, the +relative proportion, and the direction of the other organs of the flower +are altered likewise.[562] Not only is the whole symmetry changed, but +the physiological operations carried on in the flower undergo +corresponding alterations. + +There are certain co-relations which do not appear to have hitherto +attracted the attention they merit; such, for instance, is that which +exists between the particular period at which an organ is developed and +its position and form. In normal morphology this has, to some extent, +been worked out, as in the case of definite and indefinite, centrifugal +and centripetal inflorescences, and in the definite or indefinite +formation of shoots, &c. + +Other instances may be cited in the frequent co-existence of regular +flowers and definite inflorescence, the terminal position of many +peloriated flowers, the relationship between indefinite inflorescence +and prolongation of the axis, &c. + +Again, the simultaneous evolution of the parts of the flower and their +consequent verticillate arrangement, are often associated with the +production of different forms from those characteristic of organs +developed in succession, and, in consequence, arranged spirally. In the +case of simultaneous development we meet with a repetition of whorls, +as in what are termed hose-in-hose flowers (flores duplicati, +triplicati, &c.), and also with cases of peloria. In instances where the +organs are formed successively in spiral order, we meet with such +changes as median prolification, petalody, and phyllody. All these are +alterations which we might anticipate from the activity of the growing +point being checked at a certain stage in the one case, while it is +continuous in the other. This relationship between the definite and +indefinite modes of growth and the form of the several organs of the +flower, is more constant in reality than it may appear to be from a +perusal of the lists of genera in the foregoing pages, in which it was +not possible to show sufficiently well the comparative frequency of any +given changes in individual plants. Had it been possible to give +statistics setting forth the frequency of certain deviations in plants +or groups having a particular organisation, as compared with the rarity +of their occurrence in other plants of a different conformation, these +co-relationships would have been rendered much more evident. A hundred +different plants, for instance, may be named in any particular list, of +which fifty shall be of one type of structure, and the remainder of +another. And the co-relative changes in each fifty may appear to be +evenly balanced, but so far is this from being the case, that the +frequency of the occurrence of a particular change, in one species in +the list, may be so great as far to exceed the instances of its +manifestation in all the rest put together. This difficulty is only very +partially obviated by the addition of the * to signify especial +frequency of occurrence of any given malformation in the plants to whose +names it is affixed. + +=Compensation.=--But little further need be said on this head. An +atrophied condition of one part is generally associated with an +hypertrophied condition of another, and scarcely a change takes place in +one direction, but it is associated with an inverse alteration in some +other. This principle is not universal, and its application must not be +unduly strained. It requires specially to be considered in reference to +differences in the degree or kind of functional activity exercised by +the organs implicated--points beyond the scope of the present volume. + +=Teratology and classification.=--Lastly, there remain to be mentioned +the bearings of teratology on systematic botany. There are those who +would entirely exclude teratology from such matters. It may be expedient +to do so when the object sought is one of convenience and facility of +determination only, but when broader considerations are concerned, +teratology must no more be banished than variation. In most instances +the one differs but in degree from the other. If variation affords aid +in our speculations as to the affinities and genealogical descent of +species and other groups, so does teratology, and in a far higher +degree. + +Take the characters of exogens as distinct from endogens; even under +ordinary circumstances, no absolute distinction can be drawn between +them. There are plants normally of an intermediate character, while, to +take exceptional instances, there are exogens with the leaves and +flowers of endogens, and endogens whose outward organisation, at any +rate, assimilates them to exogens. Diclinous or monochlamydeous plants +owe their imperfect conformation to suppression, and may become +structurally complete by a species of peloria. Structurally +hermaphrodite flowers become unisexual by suppression, or are rendered +incomplete by the non-development of one or more of their floral whorls. +Hypogynous flowers become perigynous by adhesion, or by lack of +separation; perigynous ones become hypogynous by an early detachment +from the receptacle that bears them, or by the arrested development of +an ordinarily cup-like receptacle. + +How the relative position of the carpels and the calyx may be altered +has already been alluded to, as has also the circumstance that while it +is common to find an habitually inferior or adherent ovary becoming +superior or free, it is much more rare to find the superior ovary +adherent to the receptacle or to the calyx.[563] Regular and irregular +peloria, too, serve to show how slight are the boundaries, not only +between different genera, but also between different families. + +While, therefore, teratology may be an unsafe guide in strictly +artificial schemes, it is obvious that its teachings should have great +weight in all philosophical systems of classification. + +The questions will constantly arise, does such and such a form represent +the ancestral condition of certain plants? Is it a reversion to that +form? or is it, on the other hand, the starting point of new forms? + +Such questions cannot receive at present any satisfactory answer, but +the evidence we have seems to indicate that pre-existing forms were +simpler, and less specialised in structure than those now existing, and +hence if we meet with malformations of a simple kind, we may consider +them as possible reversions; while, if they present features of +increased complexity, and more sharply defined differentiation, we may +assume them to be evidences of a progressive rather than of a +retrogressive tendency. + +That monstrosities so called may become the starting points of new forms +is proved by circumstance that, in many cases, the peculiarities are +inherited so that a new "race" is produced and perpetuated: and if a new +race, why not a new species? The difference is one of degree only. + +FOOTNOTES: + +[553] See Clos., 'Bull. Soc. Bot. Fr.,' 1856, vol. iii, p. 679. + +[554] 'Theorie de la Feuille,' p. 26. + +[555] An additional illustration of this may be cited, which has been +brought under the notice of the writer by Dr. Welwitsch recently, and in +which some of the leaflets of the pinnate leaf of a species of +_Macrolobium_ were absent, and their place supplied by flowers arranged +in cymes. + +[556] The presence of a bud at the extremity once considered to be an +absolute distinction between branch and leaf, which latter never forms a +bud exactly at the apex--is invalidated by the case of the Nepaul +barley, p. 174. + +[557] 'Journ. Linn. Soc.,' vol. x, p. 103 _et seq._ + +[558] See also the receptacular tube (ovary?) of _Baeckea_ bearing +stamens, see p. 183. It would be natural to see stamens springing from +the receptacle but not from the ovary. + +[559] In _Passiflora_ the organogeny of the flower clearly shows the +truth of this assertion, as was indeed shown by Payer and Schleiden. + +[560] See Payer, 'Organ. Veget.' + +[561] It must, however, be borne in mind that no true leaf-organ has yet +been seen with a bud at its exact apex (unless it be the nepaul barley), +while in the case of an axial organ such a position of the bud is +constant. The nearest approach is in the case of impari-pinnate leaves +in which the terminal leaflet is jointed to the common rachis, and in +the leaves of some _Meliaceae_ which continue to push forth new leaflets +even after the leaf has attained maturity. + +[562] A singular instance of co-relation was shown by Mr. Saunders at +the Scientific Committee of the Royal Horticultural Society, February +16th, 1868, in a hyacinth with perfectly green, long, tubular, erect, +not horizontally spreading flowers. + +[563] An illustration of this latter nature in the case of a cherry, +which was surmounted by the calyx lobes, precisely as in the case of a +pomaceous fruit, has been given at p. 424, _adnot._ + + + + +APPENDIX + +DOUBLE FLOWERS.[564] + + +In ordinary language, the epithet double flowers is applied to flowers +of very varied structural conformation. The most common conditions +rendering a flower double, in the popular acceptation of the term, are +substitutions of petals or petal-like bodies for stamens and pistils, +one or both. (See Petalody, p. 283.) Another very common mode of +doubling is brought about by a real or apparent augmentation in the +number of petals, as by multiplication, fission, or chorisis. (See pp. +66, 343, 371, 376.) Sometimes even the receptacle of the flower within +the outer corolla, divides, each subdivision becoming the centre of a +new series of petals, as in some very luxuriant camellias and anemones. +The isolation of organs which, under ordinary circumstances, are united +together, is another circumstance, giving rise, in popular parlance, to +the use of the term double flower. (See Adesmy, Solution, pp. 58, 76, +82.) Prolification is another very frequent occurrence in the case of +these flowers, while still other forms arise from laciniation of the +petals, or from the formation of excrescences from the petals or +stamens, in the form of supplementary petal-like lobes. (See Enation, p. +443.) + +As these matters are all treated of under their respective headings, it +is not necessary to allude to them again in detail. It may be well, +however, to allude, in general terms, to the causes which have been +assigned by various writers for their formation, and to the means which +have been adopted by practical experimenters to secure the production of +the flowers often so much esteemed by the florist. It must be admitted +that, in spite of all that has been written on the subject, but very +little is known about these matters. In the case of the stock the +following means have been adopted by cultivators in order to obtain +plants bearing double instead of single flowers. There is first the +crossing of single flowers with double ones, effected by planting a +double-flowered plant in proximity to a single-flowered one; but this, +it is obvious, could lead to no important results, since the double +flowers, having no pollen, could not possibly influence the seed, which +is borne only by the single-flowered plants. Another plan is the +degustation of the buds, that is to say, the chewing of the well-formed +buds; it is held that the single plants can be recognised by their +sweeter taste and greater consistence, and may thus be weeded out; but +there is at least the disadvantage attending this method, that the +plants, single as well as double, must all be grown up to the period +when these buds are tolerably well advanced. A third method which has +been adopted is, that of sowing the seeds at a particular lunar epoch, +great confidence being placed in the plan of planting them during the +last quarter of the moon, but such confidence is found to be misplaced. +The plan of removing the stamens has had its supporters, but as this +must be done at an early stage of development, and could only influence +the result by diverting the vital force which would be expended in the +maturation of the pollen, to the perfecting of the seeds, it is obvious +that the plan is impracticable for all ordinary purposes, even if in any +degree efficient, which from the plasticity of vegetable development, +and the faculty of doubling which is inherent in the stock family, is +not at all improbable. Still another mark, the presence of a fifth petal +in the single or seed-bearing flower, has been held to indicate the +assurance of obtaining a crop of double-flowered plants from seeds saved +from flowers possessing this peculiarity. To a certain extent, +doubtless, this expectation would be realised, owing to the plasticity +and inherent quality just alluded to, but the proportion would be too +small for any useful practical purpose. + +"The gardeners of Erfurt," observes M. Chate, who has written a +book[565] on the subject, in which he makes known a means of obtaining +double-flowered stocks founded on more than fifty years' practice in his +family, "have, for a long time, to a certain extent monopolised the sale +of seeds of these plants. To obtain these seeds, the Erfurt gardeners +cultivate the flowers in pots, and place them on shelves in large +greenhouses, giving them only sufficient water to prevent them from +dying. So cultivated the plants become weakened, the pods shortened, and +the seeds less numerous, and better ripened; and these seeds give from +60 to 70 per cent. of double flowers. + +"The seeds from these plants are said to be mostly of an abnormal shape, +which is so striking that experienced cultivators are able to separate +those which would furnish double flowers from those which would produce +single ones." + +M. Chate's method, which he calls the French one, gives still greater +results, viz.: 80 per cent. of double flowers, and these produced by +very simple means. "When my seeds," he observes, "have been chosen with +care, I plant them, in the month of April, in good dry mould, in a +position exposed to the morning sun, this position being the most +favourable. At the time of flowering I nip off some of the flowering +branches, and leave only ten or twelve pods on the secondary branches, +taking care to remove all the small weak branches which shoot at this +time. I leave none but the principal and the secondary branches to bear +the pods. All the sap is employed in nourishing the seeds thus borne, +which give a result of 80 per cent. of double flowers. The pods under +this management are thicker, and their maturation is more perfect. At +the time of extracting the seeds the upper portion of the pod is +separated and placed aside, because it has been ascertained that the +plants coming from the seeds situated in this portion of the pod, give +80 per cent. of single flowers. They yield, however, greater variety +than the others. This plan of suppressing that part of the pod which +yields single flowers in the largest proportion, greatly facilitates the +recognition of the single-flowered plants, because there remains to be +eliminated from among the seedlings only from 10 to 15 per cent. + +This separation of the single from the double-flowered plants, M. Chate +tells us is not so difficult as might be supposed. The single stocks, he +explains, have deep green leaves (glabrous in certain species), rounded +at the top, the heart being in the form of a shuttlecock, and the plant +stout and thickset in its general aspect, while the plants yielding +double flowers have very long leaves of a light green colour, hairy, and +curled at the edges, the heart consisting of whitish leaves, curved so +that they enclose it completely. Such is the substance of M. Chate's +method of securing so large a proportion of double-flowered plants, and +then of separating them from the remaining single ones--a method which +commends itself to the good sense of the intelligent cultivator."[566] + +Signor Rigamonti, a great cultivator of pinks, asserted that he was able +to distinguish double from single-flowered pinks, in the seedling state. +According to this gentleman, those seedlings which produce three +cotyledons in a whorl in place of two, form double flowers. In the case +of _Primula sinensis_ the same results occurred. Some had three leaves +in a ring, others two; most had the leaves standing one over the other +as usual. These were divided into three sets, and when they flowered, +the first lot were all double, the second semi-double, the third single. +But these statements have not been confirmed by other observers; and the +writer can safely assert that seedling pinks occasionally produce three +cotyledons, and subsequently single flowers. He has never observed a +double flower under these circumstances, though it is true his +experience in this matter has been but small. + +A writer in Otto's 'Gartenzeitung,' considers that double flowers are a +consequence of dryness of soil and atmosphere, and not of a luxurious +soil, rich in nutritious matter, having arrived at this conclusion from +an observation of the following circumstances: + +"Fifty years ago we saw _Kerria japonica_ in a hothouse with single +flowers. Twenty years later we met with it in several gardens, in the +open air, but always with double flowers. At this time we were assured +that single-flowered plants were no more to be found in the whole of +Europe, and botanists forming herbaria offered considerable sums for a +branch of _K. japonica_ with single flowers. We were requested to take +the plant in hand for the purpose of inducing it to produce single +flowers. We were advised to plant it out in a rich soil, which was done, +but, by chance, the situation was sloping, consequently it did not +retain moisture, and all the flowers produced for several years in +succession were double. Shortly after, the captain of an English ship +again brought plants bearing normal flowers from Japan, which were soon +spread over the continent, and of which we received one plant. After +three years all the young plants raised from cuttings were +double-flowered. + +"In the year 1820 we several times visited a garden in the neighbourhood +of Vienna, well known on account of its plant culture. The gardener +there possessed an immense plant of _Camellia japonica_ with single +flowers, and some small plants raised from this by cuttings, but no +other variety of camellia. He fertilised the flowers with their own +pollen, harvested seeds, which he sowed, and the plants raised from them +were placed in an extremely dry, lofty conservatory, where, after some +years, instead of producing single flowers, they all produced double +ones. The seedlings and mother plant were planted in one and the same +kind of earth, and some of the flowers on the old plant also showed an +inclination to become double. + +"This, at that time, to us, enigmatical phenomenon, was kept in mind +until we had an opportunity of instituting comparisons between the +climate of Japan and China and our own, and we then concluded that in +the case of a plant imported from thence, and exposed to such different +climatical influences, the origin of the greater or less imperfection of +its sexual organs was probably owing to this change, as we had +experienced in _Kerria_ and _Camellia_; and that the sterility of many +other exotic plants might be attributed to the same cause. The +difference in the climatical relations of Japan and Europe is very +considerable. In Japan, previous to the new growth of _Kerria_ and +_Camellia_, a rainy season of three months' duration prevails; in +Europe, on the contrary, dry winds prevail especially in the eastern +part, where our plains are often transformed into deserts. Is it, +therefore, remarkable that a plant introduced from Japan into Europe, +exposed to the influences of this great diversity of climate, should +produce imperfect sexual organs incapable of further propagating the +plant from seeds? A rich soil, with the necessary amount of moisture, +will never engender double flowers."[567] + +Mr. Darwin[568] describes a peculiar form of _Gentiana Amarella_, in +which the parts of the flower were more or less replaced by compact +aggregations of purple scales in great numbers. A similar condition is, +indeed, not uncommon in this plant, and, as Mr. Darwin also remarked, on +hard, dry, bare, chalky banks, thus bearing out the views expressed by +the writer in the 'Gartenzeitung' just cited. Some double flowers of +_Potentilla reptans_ found growing wild near York, and transmitted to +the writer by a correspondent, were observed growing along a high wall, +in a dry border, close to a beaten path, bordering on a gravel pit, +others were found on a raised bank, which, from its elevation and +exposure to the sun, was particularly dry. + +On the other hand, the double-flowered _Cardamine pratensis_, which is +occasionally found in a wild state, always grows in very wet places. + +Of late years a remarkable double-flowered race of _Primula sinensis_ +has been obtained. In particular, Messrs. Windebank and Kingsbury, of +Southampton, have succeeded in raising a set of plants in which the +flowers are very double and very attractive in a florist's point of +view. The corollas in these flowers are not merely duplicated, but from +their inner surface spring, in some cases, funnel-shaped or tubular +petals (p. 315), so regular in form as quite to resemble a perfect +corolla. These tubes are attached to the inner side of the tube of the +corolla, in the same way as are the stamens, these latter organs being, +it appears, absent. The carpels are present, but open at the top, and +bear numerous ovules, hence it was at first surmised that these plants +were obtained and perpetuated, by the application of pollen from single +flowers to these double-flowered varieties. + +The raisers of this fine race however assert that "the double kinds are +all raised from the seed obtained from _single_ flowers; the double +blooms do not produce seed, as a rule, and even if they did yield seed, +and it were to germinate, the plants so raised would simply produce +single flowers." Semi-double flowers will produce seed, but it is +necessary that they should be fertilised with the pollen from the single +blooms. They rarely, however, if ever, produce really double flowers +when so fertilised, and the number of semi-double flowers, even, is +always small, the remainder, and, consequently, the larger part, proving +single. To obtain double varieties, the raiser fertilises certain fine +and striking single flowers, with the pollen of other equally fine +single blooms, and the desired result is obtained. This is Messrs. +Windebank and Kingsbury's _modus operandi_, the exact process or mode of +accomplishment being, however, a professional secret.[569] + +From what has been said, as well as from other evidence which it is not +necessary to detail in this place, it may be seen that the causes +assigned by physiologists, and the plans proposed by cultivators for the +production of double flowers, are reducible to three heads, which may be +classed under Plethora, Starvation, and Sterility. These three seem +inconsistent one with the other, but are not so much so as they at first +sight appear to be. + +Tho advocates of the plethora theory have much in their favour: for +instance, the greater frequency of double flowers among cultivated +plants than among wild ones. The great preponderance of double flowers +in plants derived from the northern hemisphere, when contrasted with +those procured from the southern, as alluded to by Dr. Seemann, seems +also to point to the effect of cultivation in producing these flowers. +Now, although this is, to a large extent, due to the selection that has +been for so long a period practised by gardeners, still that process +will not account for the appearance of double flowers where no such +selection has been exercised; as in the case of wild plants. Some double +peas, observed by Mr. Laxton, appeared suddenly; they had not been +selected or sought for, but they were produced, as it would appear, as a +result of high cultivation, and during the period when the plant was in +greatest vigour; and as the energies of the plant failed, so the +tendency to produce double flowers ceased. Indeed, in reference to this +subject, it is always important to bear in mind the time at which double +flowers are produced; thus, an annual plant subjected to cultivation, +will, it may be, produce single flowers for the firet year or two, then +a few partially double flowers are formed, and from these, by careful +selection and breeding, a double-flowered race may be secured. +Sometimes, as in the peas before alluded to, in the same season the +earlier blossoms are single, while later in the year double blossoms are +produced. This happens, not only in annuals, but also in perennials, and +is not infrequent in the apple; an illustration of this occurrence in +this tree is given in the 'Gardeners' Chronicle' for 1865, p. 554.[570] +Sometimes the flowers on a particular branch are double, while those on +the rest of the plant are single.[571] On these points, the evidence +furnished by a double white hawthorn in the Royal Botanic Gardens at +Edinburgh is important. Professor Balfour kindly wrote as follows in +reply to an inquiry respecting this plant:--"A double white hawthorn in +the Royal Botanic Gardens produced double flowers in spring. It retained +its leaves during autumn and winter, until the following spring. It then +flowered in the second spring, but produced weak single flowers only, +and has continued to do so ever since. The flowering has been always +weak, since this change of flowers from double to single. Mr. M'Nab +attributes the change in the duration of the leaves to the filling up of +the ground round the tree, to the height of a foot and a half on the +stem. He is now trying the effect of extra manure in giving extra vigour +to the plant." Here, at least, the production of single flowers would +seem to be the result of debilitating causes, connected with the unusual +persistence of the leaves, &c., for while the tree was healthy, double +flowers were produced. + +A similar illustration came under the writer's own notice. Some seedling +balsams, of a strain which from long selection and hereditary tendency +produces, year after year, double flowers were, in the spring (of 1866), +allowed to remain in the seed-pans for many weeks after they were ready +to be potted off; they were hence partly starved, and when they bloomed, +they produced single flowers only. But these same plants, when more +liberally treated, produced an abundance of double flowers. Moreover, +other seedlings of the same batch, but sown later, and potted off at the +usual time, produced double flowers as usual. Of a like character is the +fact that the double _Ranunculus asiaticus_ loses its doubleness if the +roots are planted in a poor soil. + +On the other hand, the way in which double stocks are stated to be +produced at Erfurt, viz.: by giving the plants a minimum supply of +water, and the other circumstances alluded to as showing the connection +between the production of double flowers, and a deficiency of water, as +well as the experiments of Mr. Monro, go to show that, so far from +plethora, the inducing cause must be more nearly allied to inanition, +though the impoverishing process is, to a certain extent, counteracted +by only allowing a few of the seed-pods to ripen, and thus concentrating +in a small number of flowers the nutriment intended for many. + +Professor Edward Morren ('Bull. Acad. Roy. Belg.,' 2me ser., vol. xix, +p. 224) considers the existence of true variegation in leaves, and the +production of double flowers, as antagonistic one to the other; the +former is a sign of weakness, the latter of strength. But it would seem +that the exceptions are so numerous--so many cases of the co-existence +of variegated leaves, and double flowers are known, at least in +individual plants if not in species--that no safe inferences can be +drawn as to this point. Since the above remarks were printed, Professor +Morren has published a second paper on the subject, upholding his former +views as to the incompatibility of variegated foliage (not mere +colouration) and double flowers. In this paper he criticises the +objections raised by the present writer and others, and examines some +of the alleged exceptions. Some of these the Belgian savant finds to +prove his rule, inasmuch as although there is a co-existence of +variegated foliage and double flowers in these illustrations, yet the +plants are weakly, the flowers ill formed, or fall off before expansion. +Admitting all this, there still remain cases in which double flowers and +variegated foliage do exist in conjunction, and where the plants are +vigorous and the flowers well developed. Instances of this are known to +cultivators in species of _Dianthus_, _Hemerocallis_, _Althaea_, +_Paeonia_, _Rosa_, _Ranunculus_, _Serissa_, _Saponaria_, etc., and +probably the art of the cultivator would speedily be successful in +raising other examples, were it a matter of importance or interest to +them to do so. At any rate, the existence of a few unimpeachable +illustrations is sufficient to support the opinion of the present +writer, and objected to so strongly by M. Morren that, in the present +state of our knowledge, "no safe inferences can be drawn" from the facts +alluded to by the Belgian professor.[572] + +Mr. Darwin[573] has thrown out the suggestion that the cause for the +appearance of double flowers may be sought for in some previous state of +things, bringing about sterility or imperfect formation, or functional +activity of the genitalia of the flower, and consequent compensatory +increase of the petaline element, either in the form of an increased +number of bracts, petals, &c., or in the substitution of petals for +stamens and pistils, &c. + +In considering these points the question arises whether they can be +reconciled one with another. And there is little doubt but that they may +be. The production of a flower is preceded by an arrest of vegetation; +this is obvious: the current of the plant's life becomes changed, the +growth of the leaves is checked, the lengthening of the branches is +arrested as the flower-bud forms; moreover, there is a close +relationship in a large majority of flowers between the outer envelopes +of the flower and the scales of a leaf-bud; this is especially so in +regard to the venation, and is admitted by all morphologists. So far, +then, it may be said that the production of a flower, like that of a +bud, is due to a diminution of vegetative action; and as in double +flowers we have, for the most part, merely a repetition and exuberant +formation of floral envelopes, so we may attribute their formation to a +continuance of the same feeble vegetative action as that which produced +the first or normal series. How, then, can a copious supply of rich +food, such as is provided by cultivation, produce double flowers? To +this question, according to our theory, the reply would be that the +quantity of food is excessive, more than the plant can properly digest; +and hence vegetative action is stopped, at least partially--pretty much +as it would be if the plant were placed in the opposite condition of +starvation. The effect of supplying a plant (or an animal) with an +excessive supply of food, which it cannot assimilate, is in many +respects similar to that which results from partially cutting off the +supplies. And the same reasoning applies to sterility. If by high +culture, or the supply of an undue quantity of nourishment, the +constitution of the plant be impaired, or if the plant be pampered, it +is no wonderful thing that sterility should ensue. Hence, then, may it +not be asserted as a general principle that in the production of double +flowers a partial arrest of development, if not of growth, however +produced, is an essential preliminary? All the attendant phenomena, such +as the obliteration of the stamens, the augmentation in the number of +floral whorls, the occurrence of prolification, are consistent with the +supposition of a primary arrest of development, more or less complete, +as the case may be: at one time permanent, at another time relaxed and +intermittent, or in a third set of cases the vegetative activity or +power of growth may be restored, and from the centre of the flower may +spring a perfect branch with perfect leaves, the production of sheaths +only being superseded by the development of leaves, in which all the +parts--sheath, stalk, and blade--are present. + +When once the disposition to form double flowers is established, that +tendency becomes hereditary: there are races of single Stocks in which, +out of hundreds of plants, scarcely one double-flowered form is met +with; but when the tendency to produce double blooms is set up, single +flowers become the exception: thus, in the Balsams, before mentioned, +not one in fifty now produces single flowers, and the seeds of these +double Balsams produce double-flowered seedlings, with scarcely a +"rogue" among them. + +The following list of plants producing double flowers of any kind is +taken from that given in 'Seemann's Journal of Botany,' vol. ii, p. 177, +and to which some additions have been made. Miscalled double flowers, +such as those of the _Compositae_, _Viburnum Hydrangea_, &c., are +excluded. + +RANUNCULACEAE. + +Clematis Viticella, _Linn._, S. Europe. + florida, _Thunb._, Japan. + Fortunei, _Moore_, Japan. + patens, _Desne_, Japan. +Anemone japonica, _Sieb. et Zucc._, Japan. + coronaria, _Linn._, S. Europe, Asia Minor. + hortensis, var. _Linn._, S. Europe. + palmata, _Linn._, N. Africa, Spain, Portugal. + nemorosa, _Linn._, Europe, N. America, Siberia. + sylvestris, _Linn._, S. Europe, Siberia. +Hepatica triloba, _Chaix._, Europe. +Ranunculus bulbosus, _Linn._, Europe, N. Amer. + repens, _Linn._, Europe, Siberia, N. Amer. + acris, _Linn._, Europe, Siberia. + aconitifolius, _Linn._, Europe. + gramineus, _Linn._, Italy, France, Portugal, Switzerland. + bullatus, _Linn._, S. Europe. + asiaticus, _Linn._, The East. +Ficaria ranunculoides, _Moench._, Europe. +Thalictrum anemoides, _Michae._, N. America. +Caltha palustris, _Linn._, Europe, Asia, N. America. +Trollius europaeus, _Linn._, Europe. + nepalensis, Himalaya. +Nigella damascena, _Linn._, Mediterranean. +Aquilegia vulgaris, _Linn._, Europe. + canadensis, _Linn._, N. America. +Delphinium Ajacis, _Linn._, S. Europe. + grandiflorum, _Linn._, Siberia, N. America. + Consolida, _Linn._, Europe, N. America. + cheilanthum, _Fisch._, Siberia. + elegans, _D. C._, North America. +Adonis autumnalis, _Linn._, Europe. + vernalis, _Linn._, Europe, Asia. +Paeonia Moutan, _Sims_, China, Japan. + officinalis, _Retz._, Europe. + tenuifolia, _Linn._, Tauria. + albiflora, _Pall._, Siberia. + paradoxa, _Andr._, S. Europe. + +NYMPHAEACEAE. + +Nelumbium speciosum, _Willd._, Africa, Asia. + +BERBERIDACEAE. + +Berberis, _sp. cult._ + +PAPAVERACEAE. + +Papaver Rhoeas, _Linn._, Europe. + bracteatum, _Lindl._, Russia. + somniferum, _Linn._, S. Europe, Asia Minor, Egypt. +Chelidonium majus, _Linn._, Europe, Asia. +Sanguinaria canadensis, _Linn._, N. America. +Podophyllum peltatum, _Linn._, N. America. + +CRUCIFERAE. + +Mathiola incana, _R. Br._, Mediterranean. + glabrata, _D. C._ + annua, _Sweet._, South Europe, Syria. +Cheiranthus Cheiri, _Linn._, Europe. +Iberis umbellata, _Linn._, Europe. + amara, _Linn._, Europe. +Cardamine pratensis, _Linn._, Europe, Asia, Africa, America. +Hesperis matronalis, _Linn._, Europe, Siberia. +Barbarea vulgaris, _R. Br._, Europe. +Sinapis arvensis, _Linn._, Europe. +Brassica oleracea. _Linn._, Europe. + +CISTACEAE. + +Helianthemum vulgare, _Spach._, Europe, N. Africa. + +VIOLACEAE. + +Viola odorata, _Linn._, Europe, Siberia. + grandiflora, _Linn._, Europe, + tricolor, _Linn._, Europe. + +CARYOPHYLLEAE. + +Dianthus barbatus, _Linn._, France, Germany. + chinensis, _D. C._, China. + Poiretianus, _Seringe_, ? + Caryophyllus, _Linn._, France, Italy. + arboreus, _Linn._, Crete. + hybridus (_gardens_). + corymbosus, _Sibth._, Asia Minor. + plumarius, _Linn._, Europe, Siberia, N. America. + deltoides, _Linn._, Europe. +Saponaria officinalis, _Linn._, Europe. +Lychnis sylvestris, _Schkr._, Europe. + vespertina, _Linn._, Europe. + flos cuculi, _Linn._, Europe. + Viscaria, _Linn._, Europe. + chalcedonica, _Linn._, Japan, Asia Minor. +Silene inflata, _Sm._; _var._ maritima, _D. C._, Europe. + +ALSINEAE. + +Sagina procumbens, _Linn._, Europe. + +MALVACEAE. + +Hibiscus Rosa sinensis, _Linn._, E. Indies. + flavescens, _Cav._, China. + alba, _Hook._, China. + syriacus, _Linn._, Syria, Carniola. +Althaea rosea, _Cav._, Caucasus, &c. +Malva rotundifolia, _Linn._, Europe. + moschata, _D. C._, Europe. + +HIPPOCASTANEAE. + +AEsculus Hippocastanum, _Linn._, Europe, N. America. + +GERANIACEAE. + +Geranium pratense, _Linn._, Europe, Siberia. + sylvaticum. _Linn._, Europe. +Pelargonium zonale, _Willd._, S. Africa. +Tropaeolum majus, _Linn._, Peru. + minus, _Linn._, Peru. +Oxalis cernua, _Thunb._, S. Africa. +Impatiens Balsamina, _Linn._, E. Ind. + +TERNSTROEMIACEAE. + +Camellia reticulata, _Lindl._, China. + Sasanqua, _Thunb._, China. + japonica, _Linn._, Japan. +Thea maliflora, _Seem._, Japan. + +AURANTIACEAE. + +Citrus Aurantium, _Linn._, Asia, South Europe. + +PAPILIONACEAE. + +Trifolium repens, _Linn._, Europe, S. America. +Medicago sp., ?., Europe. +Ulex europaeus, _Link._, Europe. +Spartianthus junceus, _Linn._, S. Europe. +Clitoria Ternatea, _Linn._, E. India. +Orobus viscoides, _D. C._, Croatia, &c. + vernus, _Linn._, Europe. +Genista tinctoria, _Linn._, Europe. + sibirica, _Linn._, Siberia. + scoparia, _Lam._, Europe. +Cytisus albus, _Link._, Portugal. +Anthyllis Vulneraria, _Linn._, Europe. +Coronilla Emerus, _D. C._, Europe. +Lotus corniculatus, _Linn._, Europe. + +ROSACEAE. + +Rosa lutea, _Mill._, Europe. + cinnamomea, _Linn._, Europe, N. America. + spinosissima, _Linn._, Central Asia. + Carolina, _Linn._, N. America. + villosa, _Linn._, Europe, Central Asia. + centifolia, _Linn._ + damascena, _Linn._, Syria. + rubiginosa, _Linn._, Europe, Asia, N. America. + moschata, _Ait._, Madeira, N. Africa. + canina, _Linn._, Europe. + alba, _Linn._, Europe, Caucasus. + indica, _Linn._, China. + nivea, _D. C._, China. + Eglanteria, _Linn._, Europe. + gallica, _Linn._, Europe, Caucasus. + pimpinellifolia, _Linn._, Europe, Central Asia. + Banksiae, _R. Br._, China. + sulphurea, _Ait._, East. +Rubus fruticosus, _Linn._, Europe. + rosifolius, _Linn._, Mauritius, E. India. + corylifolius, _Smith_, Europe. + caesius, _Linn._, Europe. +Kerria japonica, _D. C._, Japan. +Spiraea Filipendula, _Linn._, Europe. + Ulmaria, _Linn._, Europe. + prunifolia, _Sieb. et Zucc._, Japan. + Reevesii, _Lindl._, China. + strobilacea, _Sieb. et Zucc._, Japan. +Fragaria vesca, _Linn._, Europe, N. America. +Potentilla alpestris, _Hall. f._, Europe. + reptans, _Linn._, Europe, Asia. + Tormentilla, _Schrank_, Europe, Asia. + anserina, _Linn._, Europe. +Geum rivale, _Linn._, Europe. + +POMACEAE. + +Crataegus Oxyacantha, _Linn._, Europe. + Crus galli, _Linn._, N. America. +Cydonia japonica, _Pers._, Japan. +Pyrus communis, _Linn._, Europe. + Malus, _Linn._, Europe. +Eriobotrya japonica, _Lindl._, Japan. + +AMYGDALEAE. + +Amygdalus Persica, _Linn._, Persia. + communis, _Linn._, Mauritania. +Prunus domestica, _Linn._, Europe. + spinosa, _Linn._, Europe, N. America. + avium, _Linn._, Europe. + Cerasus, _Linn._, Europe. + Kerii, _Steud._, Japan. + japonica, _Thunb._, China, Japan. + insititia, _Linn._, Europe. + triloba, _Lindl._, China. + +MYRTACEAE. + +Myrtus communis, _Linn._, S. Europe. +Punica Granatum, _Linn._, S. Europe, Marocco. + +PHILADELPHACEAE. + +Philadelphus Coronarius, _linn._, S. Europe. +Deutzia Crenata, _sieb. Et Zucc._, Japan. + +ONAGRACEAE. + +Fuchsia globosa, _Lindl._ (and var. hort. pl.), Mexico. +Epilobium tetragonum, _D.C._, Europe. +Clarkia pulchella, _Pursh._, California. + elegans, _Douglas_, N. America. + +PORTULACACEAE. + +Portulaca grandiflora, _Hook_, Chili. + +GROSSULARIACEAE. + +Ribes sanguineum, _Pursh._, N. America. + +SAXIFRAGACEAE. + +Saxifraga granulata, _Linn._, Europe. + +UMBELLIFERAE. + +Daucus Carota, _Linn._, Europe. + +RUBIACEAE. + +Ixora grandiflora, _De Cand._, E. India. +Serissa foetida, _Comm._, China, Japan. +Gardenia Fortuniana, _Hook._, China. + florida, _Linn._, China, E. India. + radicans, _Thunb._, Japan. + +CAPRIFOLIACEAE. + +Lonicera Periclymenum, _Linn._, Europe. +Sambucus nigra, _Linn._, Europe. + +CAMPANULACEAE. + +Campanula latifolia, _Linn._, Europe, Asia. + Tenorei, _Morett_, Naples. + Trachelium, _Linn._, Europe. + Vidallii, _H. C. Wats._, Europe. + pyramidalis, _Linn._, S. Europe. + rotundifolia, _Linn._, Europe, N. America. + persicifolia, _Linn._, Europe. + glomerata, _Linn._, Europe, Asia. + Medium, _Linn._, Europe. + rhomboidea, _Linn._, Europe. +Platycodon grandiflorum, _D. C._, Siberia. + +ERICACEAE. + +Calluna vulgaris, _Linn._, Europe, N. America. +Rhododendron indicum, _Sweet._, E. India. + ponticum, _Linn._, Asia Minor. +Azalea nudiflora, _Linn._, N. America. + glauca, _Lam._, N. America. +Arbutus Unedo, _Linn._, S. Europe. +Erica Tetralix, _Linn._, Europe. + cinerea, _Linn._, Europe. + hyemalis, gardens. + +EPACRIDACEAE. + +Epacris impressa, _R. Br._, Australia. + +PRIMULACEAE. + +Primula villosa, _Jacq._, Europe. + Auricula, _Linn._, Europe. + denticulata, _Smith_, E. India. + acaulis, _Jacq._, Europe. + clatior, _Jacq._, Europe. + praenitens, _Ker._ = sinensis, _Lindl._, China. +Lysimachia Nummularia, _Roem et Schult._, Europe. +Anagallis tenella, _Linn._, Europe. + +JASMINACEAE. + +Jasminum officinale, _Linn._, S. Europe. + Sambac., _Ait._, E. India. + hirsutum, _Hook._, China. + grandiflorum, _Lindl._, S. Europe. + +OLEACEAE. + +Syringa persica, _Linn._, Persia. + vulgaris, _Linn._, Europe, Persia. + +APOCYNEAE. + +Vinca minor, _Linn._, Europe. + major, _Linn._, Europe. +Nerium odorum, _Ait._, E. India. + Oleander, _Linn._, S. Europe. +Tabernaemontana coronaria, _Willd._, E. India. +Allamanda cathartica, _Aubl._, S. America. + +CONVOLVULACEAE. + +Calystegia sepium, _R. Br._, Europe, America, Asia. + pubescens, _Lindl._, China. +Convolvulus tricolor, _Linn._, S. Europe. +Ipomoea pandurata, _Meyer_, S. America. + +SOLANACEAE. + +Datura cornigera, _Hook._, Peru. + fastuosa, _Linn._, S. America, Egypt. + arborea, _Linn._, S. America. + chlorantha, _Hook._ + humilis, _Desf._ +Petunia nyctaginiflora, _Juss._, S. America. + violacea, _Hook_, S. America. +Solanum Dulcamara, _Linn._, Europe. + +GENTIANACEAE. + +Gentiana Amarella, _Linn._, Europe. + +OROBANCHACEAE. + +Orobanche sp. + +SCROPHULARIACEAE. + +Mimulus luteus, _Linn._, Chili. +Antirrhinum majus, _Linn._, S. Europe. +Digitalis purpurea, _Linn._, Europe. +Linaria vulgaris, _Mill._, Europe, N. America. +Veronica, sp. +Calceolaria, var. cult. + +GESNERACEAE. + +Achimenes longiflora, _D. C._, Mexico. +Gloxinia var. hort. + +VERBENACEAE. + +Clerodendron fragrans, _Willd._, Japan. +Verbena var. hort. + +NYCTAGINEAE. + +Mirabilis Jalapa, _Linn._, Trop. America. + +LAURINEAE. + +Laurus nobilis, _Linn._, S. Europe. + Sassafras, _Linn._, N. America. + +IRIDACEAE. + +Gladiolus tristis, _Linn._, Cape of Good Hope. +Crocus aureus, _Sibth_, Europe, Asia Minor. + Susianus, _Curt._, Asia Minor. + pusillus, _Tenore_, Italy. + vernus, _Smith_, S. Europe. +Iris sibirica, _Linn._, Europe. +Iris Kaempferi, _Siebold_, Japan. + +AMARYLLIDACEAE. + +Galanthus nivalis, _Linn._, Europe. +Leucoium vernum, _Linn._, Europe. +Sternbergia lutea, _Gawl._, Europe, Asia Minor. +Hippeastrum equestre, _Herb._, S. America. +Narcissus cernuus, _Salisb._, S. Europe. + Telamonius, _Schult._, Europe. + lobularis, _Schult._ + concolor, _Schult._, Portugal. + biflorus, _Curt._, Europe. + italicus, _Ker._, Italy. + incomparabilis, _Curt._, Italy. + Cypri, _Haw._, Cyprus. + Pseudo-Narcissus, _Linn._, Europe. + poeticus, _Linn._, Europe. + Jonquilla, _Linn._, S. Europe, East. + Tazetta, _Linn._, S. Europe. + poculiformis, _Salisb._, S. Europe. + +ORCHIDACEAE. + +Orchis Morio, _Linn._, Europe. + mascula, _Linn._, Europe. + pyramidalis, _Linn._, Europe. +Ophrys fucifera, _Linn._, Europe. +See also pp. 380, 509. + +HYDROCHARIDACEAE. + +Hydrocharis Morsus ranae, _Linn._, Europe. + +ASPHODELEAE. + +Asphodelus luteus, _Linn._, S. Europe. + +LILIACEAE. + +Tulipa Gesneriana, _Linn._, Asia Minor. + sylvestris, _Linn._, S. Europe. +Scilla autumnalis, _Linn._, Europe. + nutans, _Smith_, S. Europe. +Convallaria majalis, _Linn._, Europe, America. + Polygonatum, _Linn._, Europe. +Trillium grandiflorum, _Spreng._, America. +Fritillaria Meleagris, _Linn._, Europe. + imperialis, _Linn._, Persia. +Lilium Martagon, _Linn._, Europe. + candidum, _Linn._, Syria, Persia. +Hyacinthus orientalis, _Linn._, East. +Polianthes tuberosa, _Linn._, E. India. +Hemerocallis disticha, _Don._, Nepal. + Kwanso, gardens. + fulva, _Linn._, S. Europe. + +COLCHICACEAE. + +Colchicum autumnale, _Linn._, Europe. +Tofieldia calyculata, _Wahl._, Europe. + +BUTOMACEAE. + +Sagittaria latifolia, _Willd._, N. America. + sagittifolia, _Linn._, Europe, Asia, America. + +COMMELYNACEAE. + +Tradescantia virginica, _Linn._, N. America. + alba, gardens. + +FOOTNOTES: + +[564] This appendix forms a portion of a paper published in the +'Proceedings of the International Botanical Congress,' London, 1886, p. +127, and which it has been deemed advisable to reproduce with sundry +additions and modifications. + +[565] 'Traite des Giroflees,' per E. Chate. + +[566] Leading Article in the 'Gardeners' Chronicle,' p. 74, 1866. + +[567] Otto's 'Gartenzeitung,' 1866. + +[568] 'Gard. Chron.,' 1843, p. 628. + +[569] 'Gard. Chron.,' 1867, p. 381.--Art. "Chinese primroses." + +[570] See also p. 79, fig. 36. A similar flower is figured in 'Hort. +Eystett. Ic. Arb. Vern.,' fol. 5. "Fructus nondum observatus est +fortassis alimento uberius in flores refuso, nullus sperari possit." + +[571] See De Candolle, 'Plant. Rar. Genev.,' 1829, p. 91; and Alph. de +Candolle.' Geog. Bot.,' p. 1080. + +[572] See 'Gardeners' Chronicle,' 1868, p. 1113. + +[573] Ibid., 1843, p. 628. + + + + +NOTE. + + +During the progress of the foregoing pages through the press, several +additional illustrations of particular malformations have come under +notice. Some of the more important of these may here be recorded. + +_Fasciation_ (see p. 11).--The following plants may be added to the +list:--_Acer eriocarpum_, _Arabis albida_, _Brassica oleracea_, var., +_Guarea_, sp., _Artabotrys_ sp. In all, with the exception of the +first-named, the fasciation occurred in the inflorescence. In some +species of _Artabotrys_, indeed, fasciation and curvation of the +inflorescence are common. + +_Synanthy_ (p. 39).--Several additional instances of adhesion of two or +more flowers in _Calanthe vestita_, _C. Veitchii_, and other forms of +this genus may be cited. These furnish further illustrations of the much +greater liability of some plants to particular changes as compared with +others. _Scilla bifolia_, _Gagea arvensis_, and _Viola odorata_ may be +added to the list of synanthic plants. + +_Alterations of placentation, &c._ (see pp. 98, 483).--M. Casimir De +Candolle, in a letter to the author, dated March 8th, 1869, thus writes +of the existence of a double row of carpels in _Pyrus spectabilis_ and +_Crataegus Oxyacantha_, "a longitudinal section of a double flower of +_Pyrus spectabilis_ shows two rows of carpels, placed one above another. +The arrangement of the vascular bundles shows that the upper row is +external in relation to the lower series. The carpels of the latter are +wholly coalescent as in a pear, while those of the upper verticil are +only partially coherent or sometimes quite distinct. The placentation is +constantly axile in the inferior row and parietal in the upper one. The +number of ovules in each carpel of the superior row varies greatly, and +they are often, but not always, inserted in two longitudinal ranks, as +is constantly the case in the lower carpels. Double flowers of _Crataegus +Oxyacantha_ present the same anomalies." For analogous instances in +_Digitalis_, see p. 98. See also p. 380, _Saxifraga_. + +_Prolification_, p. 120.--A. P. De Candolle, "Organographie Vegetale," +tab. 40, figures an instance of suppression of one lobe of the ovary in +_Iris chinensis_, and of the presence at the base of the flower of an +adventitious and imperfect flower-bud, as in the _Phlomis_, mentioned at +p. 119. + +_Monoecious Misleto_, p. 193.--In this specimen, exhibited at one of the +meetings of the Scientific Committee of the Royal Horticultural Society +in 1869, there were both male and female flowers on the same bush. The +plant was of the male sex, with numerous long slender whip-like, +somewhat pendulous, branches bearing comparatively large broad yellowish +leaves, and fully developed male flowers at the end. From the side of +one of these male branches, near the base, protruded a tuft of short, +stiff branches, bearing small, narrow, dark green leaves, ripe berries +and immature female flowers. There was no evidence of grafting or +parasitism, of the female branch on the male, the bark and the wood +being perfectly continuous so that the only tenable supposition is that +this was a case of dimorphism. + +_Adventitious leaflet and pitcher_, see pp. 30 and 355. In a species of +_Picrasma_, in which the leaves are impari-pinnate and spread +horizontally, an adventitious leaflet was observed to project at right +angles to the plane of the primary leaf. It emerged at a point nearly +corresponding to that at which the normal pinnae were given off. The +appearance presented was thus like that of a whorl of three leaves, +except that the shining surface of the adventitious leaflet, +corresponding to the upper face of the normal leaflets, was directed +towards the axis, _i.e._, away from the corresponding portion of the +neighbouring pinnae, while the dull surface, corresponding to the lower +part of an ordinary leaflet, looked towards the apex of the main leaf, +or away from the axis. In one instance, a stalked pitcher was given off +from the same point as that from which the supernumerary leaflet +emerged, the pitcher being apparently formed from the cohesion +(congenital) of the margins of a leaflet. + +In the normal leaf of this plant there is between the bases of the +pinnae, a small reddish gland or stipel? attached to, or projecting from, +the upper surface of the rachis. It appeared from some transitional +forms that the adventitious leaflet, just mentioned, was due to the +exaggerated development of this gland, but no clue was afforded as to +the origin of the ascidium. It was not practicable to examine the +arrangement of the vascular bundles in the rachis. + +_Additional labella in Phaius._--A flower of _Phaius grandiflorus_ was +found in the same condition as the _Catasetum_, mentioned at pp. 291 and +382. + +_Tubular stem._--A species of _Sempervivum_, exhibited by Mr. Salter, of +Hammersmith, at one of the summer exhibitions of flowers at the Royal +Horticultural Society in 1868, under the name of _S. Bollei_, deserves +notice from its bearing on the question of such structures as the +calyx-tubes, the hip of the rose and such like, see pp. 394, 482. In +this plant the leaves appeared to be arranged some on the outside, +others on the inside, of an erect hollow cylinder, some six inches in +height. The oldest leaves were outside, the youngest within, so that the +appearance presented was as if the summit of the axis had been pushed or +drawn in, much as the finger of a tight glove might be invaginated in +withdrawing it from the hand. + +The plant in question thus furnishes an actual illustration of the +supposititious case mentioned at p. 482. + +_Double flowers_, see pp. 499, et seq.--The following species may be +added to those already recorded: _Lychnis coronaria_, _Hibiscus +mutabilis_, _Lotus major_, _Pisum sativum_, _Godetia_ sp., _Ipomoea +purpurea_, _Convolvulus minor_, _Heliotropium peruvianum_, _Trillium +grandiflorum_, and _Phaius grandiflorus_. + + + + +INDEX TO SUBJECTS. + + +A. + +Abortion, 455, 467 + of axile organs, 455 + calyx, 460 + corolla, 460 + indusium, 467 + leaves, 458 + ovules, 466 + perianth, 460 + pistil, 464 + receptacle, 457 + stamens, 463 + +Acaulescence, 393 + +Acaulosia, 393, 456 + +Acheilary, 398 + +Adesiny, 58, 76 + +Adhesion, 32 + of axes, 50, 55 + embryos, 56 + leaves, 33 + parts of flower, 34 + roots, 53 + +Adventitious buds, 156, 176 + flowers, 174, 176, 180 + gemmae, 173 + leaves, 162 + roots, 156 + shoots, 161 + +Albinism, 337 + +Alternation, 3, 485 + +Androecium, enlargement of, 430 + meiophylly of, 398 + meiotaxy of, 405 + of orchids, 380 + pleiotaxy of, 379 + polyphylly of, 361 + +Androgynism, 193 + +Anther (see connective), contabescence of, 463 + morphology of, 291 + ovuliferous, 200 + petalody of, 291 + sutures of, 291 + +Apilary, 397 + +Apostasis, 440 + +Aphylly, 395 + +Arrangement, 1 + +Ascidia, 30, 313 + bibliography, 30 + plants with, 30 + +Atrophy, 454 (see abortion) + +Antophyllogeny, 355 + +Avalidouires (vines), 211 + +Axes, abortion of, 455 + adhesion of, 50, 55 + cohesion of, 9 + enation from, 444 + enlargement of, 418 + fission of, 60 + suppression of, 398 + +Axile organs, see Axes + + +B. + +Barley, Nepaul, 174 + +Bigarades cornues, 303 + +Bladder-plums, 465 + +Bracts, multiplication of, 358 + staminody of, 298 + phyllody of, 242 + +Buds, adventitious on fruits, 178 + on leaves, 170, 174 + in ovary, 180 + on petals, 177 + in pith, 171 + on roots, 160 (see flower-buds, prolification) + variations of, 336 + +Bulbs, displacement of, 84 + multiplication of, 172, 350 + +Burrs, 347, 420 + + +C. + +Catacorolla, 450 + +Calycanthemy, 283 +Calyphyomy, 34 + +Calyx, abortion of, 461 + dialysis of, 70 + meiophylly of, 396 + meiotaxy of, 403 + obsolete, 460 + petalody of, 283 + pleiotaxy of, 374 + polyphylly of, 359 + solution of from ovary, 77 + +Calyx-tube, 394, 480, 509 + +Carnation wheat-ear, 371 + +Carpels (see Pistil, Fruit, Ovary), adventitious, 182 + dialysis of, 73 + enation from, 453 + fission of, 68 + in ovary, 182 + +Cauliflower, 421 + +Cenanthy, 408 + +Chloranthy, 273, 279 + bibliography, 280 + remarks on, 279 + plants subject to, 280 + +Chorisis, 59, 343 + parallel, 344 + collateral, 344 + +Chromatism, 339 + +Cladodes, 328 + +Classification, teratology in relation to, 488 + +Cohesion of leaves, 21, 25 + of petals, 28 + pistils, 29 + sepals, 27 + stamens, 29 + stems, 9 + +Colour, alterations of, 337 + +Columella, 395 + +Coniferae, leaves of, 217, 352, 484 + inflorescence of, 245 + scales of, 192, 245, 484 + +Compensation, 488 + +Connective, petalody of, 293 + +Consistence, alterations of, 432 + +Contabescence, 463 + +Contortion, 317 + +Co-relation, 486 + +Cornute leaves, 328 + +Corolla, abortion of, 461 + dialysis of, 71 + duplicate, 376 + hose in hose, 377 + meiophylly of, 397 + meiotaxy of, 403 + pleiotaxy of, 374 + polyphylly of, 359 + virescence of, 338 + +Cotyledons, increased number of, 370 + shoots, below, 161 + +Cuttings, formation of, 159 + + +D. + +Dedoublement, 59 (see Chorisis) + +Deflexion, 209 + +Deformities, 311 + +Degeneration, 470 + +Depauperate ferns, 466 + +Diadelphia, 29 + +Dialysis, 58, 69 + of calyx, 71 + carpels, 73 + corolla, 72 + plants subject to, 72 + of leaves--margins of, 70 + stamens, 73 + +Diaphysis, 103 + +Dioecious plants, 192, 193 + +Dimorphism, 333 + +Dimorphic flowers, 403 + +Direction, changes of, 201 + +Diremption, 87 + +Disjunction, see Dialysis, Fission, Solution + +Displacement, 84 + of bulbs, 84 + carpels, 96 + flower-parts of, 91 + inflorescence, 85 + leaves, 87 + ovules, 96 + placentas, 96 + +Distension, 419 + + +E. + +Ecblastesis, 107, 138 + +Elongation, 433 + of parts of flower, 438 + flower-stalks, 435 + inflorescence, 434 + leaves, 437 + nucleus of ovule, 269 + placenta, 440 + receptacle, 440 + root, 434 + thalamus, 440 + +Embryos, adhesion of, 56 + increased number of, 369 + +Enation, 443 + from axile organs, 444 + carpels, 453 + corolla, 449 + foliar organs, 445 + sepals, 448 + stamens, 453 + +Enlargement, 417 + of androecium, 430 + axile organs, 418 + buds, 420 + flower stalk, 421 + fruit, 431 + gynoecium, 430 + perianth, 428 + placenta, 425 + +Epanody, 226 + +Epistrophy, 226 + +Etiolation, 337 + +Eversion, 204 + +Excrescences, 444 (see Enation) + + +F. + +Fasciation, 11 + plants affected with, 20, 508 + +Fastigiation, 202 + +Ferns, crested, 63, 447 + depauperated, 466 + exindusiate, 467 + supra-soriferous, 189 + +Filaments, see Stamens + petaloid, 290 + 4-winged, 290 + +Finger and toe, 69 + +Fission, 59 + of carpels, 68 + leaf-organs, 61, 66 + plants, subject to, 66 + of petals, 66 + stem-organs, 60 + stamens, 68 + +Flattening, 328 + +Floral organs, displacement of, 91 + elongation of, 439 + metamorphy of, 281 + +Florets, increased number of, 351, 390 + +Flowers, adventitious, 174 + on fruits, 177 + on leaves, 174 + in ovary, 180 + on petals, 177 + on spines, 177 + apetalous, 404 + double, 490, 510 + hermaphrodite, 196 + homomorphic, 188 + increased number of, 390 + mutilated, 403 + unisexual, 193 + +Flower-stalk, enlargement of, 421 + +Flower-bud (see Prolification), replaced by leaves or scales, 164 + in place of leaf-buds, 176 + +Foliar organs (see leaves), adhesion of, 32 + +Foliar organs, cohesion of, 21, 25 + enation from, 445 + fission of, 61 + suppression of, 396 + +Form, alterations of, 213 + juvenile, persistence of, 217 + +Frondescence, 241, 279, see Phyllody, Virescence + +Fruit, adhesion of, 44 + enlargement of, 431 + + +G. + +Gemmae, formation of, 173 + +Glands, formation of, 473 + +Gnaurs, 158, 347, 417, 419 + +Grafting, 53, 56 + +Greffe des Charlatans, 56 + +Growth interrupted, 327 + irregular, 228 + +Gymnaxony, 211 + +Gynantherus, 305 + +Gynoecium (see pistils) + enlargement of, 430 + meiophylly of, 399 + meiotaxy of, 406 + pleiotaxy of, 388 + polyphylly of, 363 + suppression of, 406 + + +H. + +Hairs, formation of, 472 + +Hermaphroditism, 197 + +Heterogamy, 190 + +Heteromorphy, 311 + +Heterophylly, 330 + +Heterotaxy, 156 + +Homology, 476 + +Homomorphy, 188 + +Hose in hose corollas, 291, 377 + +Hypertrophy, 415, see Enlargement + + +I. + +Independence, 58 + +Indusium, abortion of, 467 + +Inflorescence, displacement of, 84 + elongation of, 434 + prolification of, 102, 115 + +Interrupted growth, 327 + +Inversion of organs, 206 + +Irregularity, 213 + +Irregular growth, 228 + + +K. + +Knaurs, (see gnaurs) + +Kail, 426 + + +L. + +Laciniation, see fission + +Layering, 156 + +Leaders, formation of, 203 + +Leaf-sheath, 477 + +Leaves, see foliar organs + abortion of, 458 + adhesion of, + by surfaces, 33 + to stem, 34 + adventitious, 162-165, 509 + cornute, 328 + displacement of, 86 + elongation of, 437 + enlargement of, 421 + frondiferous, 355 + geminate, 352 + multiplication of, 358 + nature of, 477 + palmate-passage of to pinnate, 439 + spiral torsion of, 326 + supernumerary, 353 + +Lily, double white, 375 + + +M. + +Meiotaxy of androecium, 405 + of calyx, 403 + corolla, 403 + gynoecium, 405 + +Meiophylly of androecium, 398 + of calyx, 397 + corolla, 397 + gynoecium, 399 + perianth, 397 + +Mellarose, 134 + +Metaphery, 91 + +Metamorphy, 239, 281 + calycanthemy, 283 + chloranthy, 273 + petalody, 283 + phyllody, 241 + pistillody, 302 + sepalody, 282 + staminody, 298 + +Mischomany, 348 + +Monadelphia, 29 + +Monoecious, 192, 193, 509 + +Monosy, 58 + +Morphology, 479 + +Multiplication, see pleiotaxy, pleiophylly + of bracts, 358, 371 + bulbs, 350 + cotyledons, 370 + embryos, 369 + florets, 351 + foliar organs, 352 + whorls, 371 + + +N. + +Nepaul Barley, 174 + +Number, alterations of, 341 + increased, 343, 353 + diminished, 392 + + +O. + +Orchids, androecium of, 380 + prolification in, 153 + +Organs, rudimentary, see Atrophy + +Ovary, inferior, nature of, 394, 482 + solution from calyx, 77 + stamens in, 184 + +Ovules, abortion of, 466 + increase of, 367 + in place of pollen, 200 + polliniferous, 183 + petalody of, 297 + phyllody of, 262 + pistillody of, 310 + suppression of, 407 + malformations of, 262 + bibliography of, 272 + + +P. + +Parasitical plants, 55 + +Peduncles, elongation of, 435 + +Peloria, 207, 228 + bibliography of, 227, 239 + regular, 219 + plants subject to, 226 + irregular, 229 + plants, subject to, 239 + +Perianth, abortion of, 460 + enlargement of, 428 + meiophylly of, 396 + pistillody of, 303 + pleiotaxy of, 375 + +Persistence, xxxvi _adnot._, 217 + +Petalody, 283 + of accessory organs, 297 + anther, 291 + calyx, 283 + connective, 292 + ovules, 297 + pistils, 297 + stamens, 284 + plants, subject to, 295 + +Petals, cohesion of, 28 + enation from, 448 + fission of, 66 + phyllody of, 251 + staminody of, 298 + tubular, 23, 314 + +Phyllode, 328 + +Phyllody, 240 (see Virescence, Chloranthy) + of accessory organs, 272 + bracts, 242 + calyx, 244 + plants subject to, 250 + in Conifers, 245 + of corolla, 251 + plants subject to, 252 + of ovules, 262 + plants subject to, 271 + pistils, 256 + plants subject to, 261 + stamens, 253 + plants subject to, 256 + +Phyllomania, 352 + +Phyllomorphy, see Phyllody + +Phyllotaxy, 1, 320 + +Pistil, abortion of, 464 + cohesion of, 29 + petalody of, 296 + staminody of, 298 + +Pistillody, 302 + of ovules, 268, 310 + perianth, 302 + sepals, 302 + stamens, 303 + plants subject to, 310 + +Pitchers, 30, 313, 509, see Ascidia + +Placentation, changes in, 96, 508 + bibliography, 100 + nature of, 483 + +Placenta, elongation of, 439 + enlargement of, 421 + +Pleiomorphy, 228 + +Pleiophylly, 353 + +Pleiotaxy, 371 + of androecium, 375 + bracts, 371 + calyx, 374 + corolla, 376 + gynoecium, 388 + perianth, 375 + +Plien, 346 + +Plymouth Strawberry, 275 + +Polyadelphia, 29 + +Pollen in ovules, 183 + abortion of, 463 + replaced by ovules, 200 + +Polyclady, 346 + +Polycotyledony, 370 + +Polyembryony, 369 + +Polymorphy, 328 + +Polyphylly of androecium, 361 + of calyx, 350 + corolla, 359 + plants subject to, 360 + of flower, 363 + gynoecium, 363 + +Polyphylly, bibliography, 364 + plants subject to, 364 + +Position, changes of, 83 + relative, 485 + +Prolification, 100 + axillary, 138 + foliar, 141 + floral, 142 + plants affected with, 148 + bibliography of, 154 + complicated, 151 + of embryo, 155 + of flower, 115 + coincident changes, 128 + median foliar, 116 + median floral, 119, 508 + plants affected with, 137 + of fruit, 134 + inflorescence, 102 + bibliography, 115 + median foliar, 103 + median floral, 105 + lateral floral, 107 + lateral foliar, 106 + + +R. + +Receptacle, abortion of, 457 + elongation of, 116, 440 + spiral torsion of, 325 + +Reflexion, 209 + +Regularity, 213 + +Rhizotaxy, 1 + +Rose Willow, 166, 168 + +Roots, adventitious, 156 + elongation of, 434 + +Rudimentary organs, 469 + + +S. + +Saint Valery Apple, 135, 282, 304, 375, 388 + +Savoys, 426 + +Scales, formation of, 164, 448, 470 + +Scape, leaves on, 163 + +Seeds, abortion of, 407 + union of, 50 + +Sepals, adhesion of, to petals, 34 + cohesion of, 27 + enation from, 448 + phyllody of, 243 + pistillody of, 303 + staminody of, 298 + +Sepalody, 282 + +Separation, 58 + +Sex, changes of, 190, 509 + +Shamrock, four-leaved, 356 + +Shoots below cotyledons, 167; _see_ Leaders + +Size, alterations in, 411 + +Solenaidie, 21, 316 + +Solution, 59, 76 + bibliography, 82 + of calyx, 77 + plants subject to, 82 + of stamens, 82 + +Spathes, increased number of, 357 + +Speiranthy, 91, 325 + +Spiral torsion, 319 + plants subject to, 325 + of leaf, 326 + of receptacle, 324 + +Spines, 456 + +Sports, 336 + +Spurs, formation of, 228, 315 + +Stamens, see Androecium + abortion of, 463 + adhesion of, 34, 35 + cohesion of, 29 + compound, 294, 345 + dialysis, 73 + enation from, 453 + fission of, 68 + in ovary, 183 + petalody of, 283 + phyllody of, 253 + pistillody of, 303 + tubular, 316 + +Staminody, of accessory organs, 301 + of bracts, 298 + petals, 298 + pistils, 299 + sepals, 298 + +Stasimorphy, 216 + +Stem, see axes + +Stipules, increased number of, 357 + +Strawberry, Plymouth, 275 + blind, 195 + +Suppression, 393 + of androecium, 405 + flower, 408 + foliar organs, 395 + +Suppression of ovules, 407 + remarks on, 409 + of seeds, 407 + +Symmetry, 213 + +Synanthy, 37 + bibliography, 45 + plants subject to, 44, 508 + +Syncarpy, 45 + +Syngenesia, 29 + +Synophty, 57 + +Synspermy, 50 + + +T. + +Tendrils, adventitious, 326 + formation of, 473 + +Thalamus, see Receptacle + +Thorns, 456 + +Torsion spiral, 319 + +Tubers, 421 + in axils of leaves, 142 + +Tubes, formation of, 312, 509 (see Ascidia, Solenaidy, Spurs) + +Tubular petals, 314 + stamens, 316 + + +U. + +Union, 8 + +Unisexuality, 195 + +Uovoli, 420 + + +V. + +Varieties, dwarf, 411 + +Venation, 338 + +Virescence, 338 + +Viviparous plants, 106, 168 + + +W. + +Warts, 444 + +Wheat-ear carnation, 371 + + + + +INDEX OF NAMES OF PLANTS. + + +[In the following Index the names of the orders that are incidentally +mentioned are printed in small capitals, those of the genera and species +in ordinary type. The names are inserted as found in the several +records, &c., without in general any attempt having been made to +determine their accuracy. For this reason the authority for the specific +name is rarely given, such citations being here unnecessary if not +impracticable. It may, however, be assumed that the names made use of +are those generally adopted by naturalists. + +This index will be found useful for statistical purposes. It will show +at a glance, at least approximately, how often certain genera and +species are affected with malformation, as contrasted with others. The +nature of the malformation may of course be ascertained by referring to +the particular page indicated by the number. The proportion of wild to +cultivated plants may also be approximately ascertained, and the effects +of cultivation estimated. The disproportionate frequency with which some +species are affected, e.g., _Trifolium repens_, &c., as contrasted with +other closely allied, and perhaps equally common species, under +apparently identical conditions, is also made manifest.] + + + +A. + +Abies Brunoniana, 245 + excelsa, 21, 61, 192, 298, 325, 456, + Larix, 90 + pectinata, 52 + +Acacia, 329 + +Aceras anthropophora, 238 + +Acer, 359, 364, 367, 508 + platanoides, 66, 459 + pseudo-platanus, 20, 371 + +Aceranthus, 225, 226 + +Achimenes, 106, 114, 296 + longiflora, 506 + +Aconitum, 148, 231, 359, 360, 399, 404, + 407, 464 + Napellus, 39, 44, 238 + +Acorus, 225 + +Actaea spicata, 251, 252 + +Adenium obesum, 417 + +Adenorophium luxurians, 254 + +Adonis, 262, 295 + autumnalis, 500 + vernalis, 500 + +Adoxa, 367 + +AErides odoratum, 398 + +AEschynanthus, 44, 297 + +AEsculus Hippocastanum, 50, 66, 295, + 369, 438, 459, 501 + +Affonsea, 364 + +Afzelia, 397 + +Agaricus, xxiii, 54 + +Agave, 177, 432 + Americana, 33, 45 + +Angelica, 365 + +Agrimonia, 137, 406 + +Agrostemma, 148 + Githago, 271 + +Agrostis alba, 169 + +Ailanthus glandulosa, 21 + +Aira alpina, 169 + caespitosa, 169 + +Ajuga Iva, 404 + pyramidalis, 20 + reptans, 243 + +Alcea, 149, 297 + +Alchemilla minima, 171 + +Aldrovanda vesiculosa, 86 + +Alisma, 115, 329 + natans, 107 + parnassifolia, 167 + +ALISMACEAE, 115 + +Allamanda, 296 + cathartica, 288, 390, 505 + +Allium, 106, 114, 170, 299, 360, 365, 367 + fragrans, 369 + vineale, 150 + +Alliaria officinalis, 269, 271 + +Almond, see _Amygdalus_. + +Alnus, 349 + fruticosa, 192 + imperialis, 459 + incana, 21 + glutinosa, 66, 244, 349 + laciniata, 65 + +Alopecurus pratensis, 169 + +Alsine media, 67, 252, 404 + +Alstroemeria, 319, 326 + +Althaea, 295 + rosea, 20, 501 + +Alyssum, 137 + incanum, 252 + +AMARYLLIDACEAE, 115, 138, 150 + +Amaryllis, 307, 310, 432 + +Ambrina ambrosioides, 397 + +AMENTACEAE, 114, 435 + +Amorpha, 21, 397 + fruticosa, 23, 30, 243 + +AMYGDALEAE, 500; see _Rosaceae_ + +Amygdalus, 122, 137, 155, 295, 297, 364 + communis, 250, 252, 369, 503 + Persica, 74, 176, 503 + +Anagallis, 73, 138, 141, 150, 288, 296, 360, 365, 367 + arvensis, 117, 161, 256, 263, 271, 278, 281, 284 + collina, 44, + phoenicea, 141, 253, 271, 441, 461 + tenella, 505 + Webbiana, 281 + +Anomodon alternatus, 49 + +Ananassa, 350 + +Anchusa, 132, 138, 339, 365 + ochroleuca, 125, 259, 262, 281 + paniculata, 262 + +Androsace maxima, 16, 20 + +Anemiopsis californica, 63, 66 + +Anemone, 113, 121, 136, 140, 142, 148, 258, 289, 295, 296, 297, 360, + 374, 430, 463, 491, (see _Hepatica_) + coronaria, 107, 250, 256, 339, 499 + hortensis, 107, 250, 339, 499 + japonica, 161, 499 + nemorosa, 250, 256, 339, 499 + palmata, 499 + +Anemone pavonina, 499 + Pulsatilla, 250 + rivularis, 165 + sylvestris, 250, 499 + +Angelica, 114, 137, 143, 150 + Razoulzii, 244, 437 + +Anthemis arvensis, 20 + nobilis, 20 + retusa, 44 + +Anthoxanthum, 61 + odoratum, 61 + +Anthriscus, 442 + +Anthurium Scherzerianum, 358 + +Anthyllis, 295 + +Antirrhinum majus, 20, 22, 24, 31, 44, 57, 73, 82, 104, 114, 121, 131, + 225, 226, 227, 230, 233, 235, 238, 253, 272, 296, 301, 315, 316, + 318, 363, 365, 378, 398, 505 + +Apargia, 114 + autumnalis, 20 + +Apium, 82, 113, 150 + graveolens, 66, 158 + Petroselinum, 370, 437 + +APOCYNACEAE, 137 + +Apple, St. Valery, 135, 282, 304, 375, 388 + +Aquilegia canadensis, 500 + Skinneri, 266, 271 + vulgaris, 24, 70, 127, 136, 220, 226, 252, 257, 260, 261, 271, 280, + 286, 287, 288, 293, 295, 74, 390, 500 + +Arabis, 148, 508 + alpina, 397, 461, 463 + pumila, 170 + sagittata, 44 + +Araucaria, 245 + +Arbutus Unedo, 291, 292, 296, 377, 504 + +Archidendron, 365 + +Ardisia serrulata, 369 + +Arenaria serpyllifolia, 461 + tetraquetra, 398, 399, 464, 469 + +Armeria, 114 + +Aristolochia Clematitis, 38, 45, 314 + caudata, 231, 238 + sipho, 23 + +Armoracia rusticana, 64, 299 + +Arnoseris, 114 + +Artabotrys, 456, 508 + +Artemisia, 405 + +Artocarpus, 407 + +Arum, 329 + maculatum, 66, 225, 227, 245, 358 + +Asparagus officinalis, 12, 19, 21, 320, 325, 471 + +Asphodelus, 138, 296 + +Asphodelus luteus, 506 + ramosus, 298, 310 + +Asplenium Trichomanes, 190 + +Astrantia, 114 + major, 368 + +Athamanta, 82, 137, 149 + Cervaria, 250, 442 + +Atriplex, 227 + hortensis, 224 + +Atropa Belladonna, 44 + +Aucuba, 21 + japonica, 21, 197 + +AURANTIACEAE, 137, 149, 502 + +Avena, 319, 351, 391 + chinensis, 298 + +Azalea, 35, 114 + glauca, 504 + indica, 44, 73, 166, 209, 289, 296 + nudiflora, 504 + + +B. + +Babingtonia, 185 + +Baeckea diosmifolia, 183 + +BALSAMINEAE, 359, 404, 501 + +Balsam, see _Impatiens_ + +Bamboo, see _Bambusa_ + +Bambusa, 307, 310, 324, 365 + +Barbarea, 295 + vulgaris, 310, 500 + +Barkhausia, taraxacifolia, 20 + +Barley, Nepaul, 174 + +Bauhinia, 328 + +BEGONIACEAE, 114 + +Begonia, 31, 81, 106, 114, 162, 170, 352 + frigida, 199, 303, 310 + fuchsioides, 281 + phyllomaniaca, 170 + +Bellevalia, 408, 461, 467 + comosa, 348 + +Bellis perennis, 17, 20, 31, 106, 114, 164, 244 + +Berberis, 272, 295, 458, 460, 500 + vulgaris, 20 + +Beta, 19, 325, 365 + vulgaris, 20 + +Betonica Alopecuros, 42, 44, 226 + +Betula alba, 66, 346, 456, 472 + populifolia, 66 + +Bidens, 114, 165, 223 + +BIGNONIACEAE, 222 + +Bignonia, 272, 296, 327 + +Bikkhia, 80 + +Blitum, 45 + polymorphum, 397, 458 + +Bocconia, 224 + cordata, 310 + +Bowiea volubilis, 409 + +BORAGINACEAE, 132, 138 + +Bouchea hyderabadensis, 166 + +Bougainvillea, 339 + +Brachythecium plumosum, 49 + +Brassica, 20, 136, 139, 295, 364, 367 + Napus, 27, 205 + oleracea, 30, 33, 66, 67, 106, 113, 136, 141, 142, 148, 250, 252, + 264, 271, 280, 351, 408, 421, 426, 442, 445, 501, 508 + Rapa, 181 + +Breynia, 198 + +Bromelia, 103 + +Bromus velutinus, 358 + +Broussonettia papyrifera, 331, 459 + +Bruniaceae, 80, 81 + +Brunia microphylla, 81 + +Bryonia, 360, 367 + +Bryophyllum calycinum, 158, 171, 483 + proliferum, 166 + +Bryum caespititium, 49 + +Bunias, 97, 136, 280, 360 + +Bunium creticum, 159 + flexuosum, 20 + +Bupleurum, 149, 459 + falcatum, 15, 20 + +BUTOMACEAE, 507 + +Buxbaumia indusiata, 49 + +Byrsonima, 137 + +BYTTNERIACEAE, 362 + + +C. + +Cabomba aquatica, 458 + +CACTACEAE, 81, 113, 149, 395 + +Cactus, 160 + +Cachrys taurica, 197 + +Caesalpinia, 365 + digyna, 48 + +Cakile maritima, 246, 250 + +Calanthe, 227, 398, 402, 508 + vestita, 39, 45, 227, 402, 508 + +Calceolaria, 41, 44, 230, 233, 284, 296, 397, 405, 406, 505 + crenatifolia, 238 + floribunda, 316 + rugosa, 238 + +Calendula, 114, 138, 339, 370 + officinalis, 252, 280, 339 + +Calla palustris, 357 + +Callitriche, 196 + autumnalis, 406 + vernalis, 406 + +Caltha, 136, 148, 295 + palustris, 141, 250, 442, 500 + +Calluna, 296, 504 + +Calycophyllum, 249, 283, 429 + +Calystegia, 114, 296 + +Calystegia Sepium, 505 + pubescens, 505 + +Camellia japonica, 288, 295, 297, 491, 494, 502 + reticulata, 502 + Sasanqua, 502 + +CAMPANULACEAE, 80, 114, 127, 131, 138, 139, 150, 334 + +Campanula, 71, 72, 73, 82, 138, 150, 250, 285, 296, 365, 367, 404, + 442, 472 + canescens, 403 + colorata, 403 + glomerata, 242, 300, 504 + latifolia, 504 + Medium, 20, 37, 44, 61, 251, 448, 504 + persicifolia, 44, 284, 300, 429, 504 + pyramidalis, 281, 504 + Rapunculus, 429 + rapunculoides, 20, 252, 300, 310, 375 + rhomboidea, 504 + rotundifolia, 377, 378, 504 + Tenorei, 504 + thyrsoidea, 20 + Trachelium, 504 + Vidallii, 504 + +Campanumaea, 80, 81 + +Camphorosma monspeliaca, 456 + +Canna, 285 + +Cannabis, 82, 194, 197 + sativa, 81 + +Cannamois virgata, 197 + +CAPPARIDACEAE, 148, 390 + +CAPRIFOLIACEAE, 45 + +Capsella bursa pastoris, 298, 361 + +Cardamine, 295, 357 + hirsuta, 398 + Impatiens, 404 + pratensis, 65, 170, 181, 495, 500 + sylvatica, 398 + +Carduus arvensis, 20 + crispus, 166, 339 + heterophyllus, 260, 262, 250 + tataricus, 250, 260, 262 + +Carex, 115, 138, 150, 191, 194, 350 + acuta, 143, 198 + caespitosa, 199 + glauca, 143, 199 + maritima, 369 + paludosa, 199 + vulpina, 428 + +Carica Papaya, 199 + +Carlemannia, 398 + +Carlina, 114 + vulgaris, 20 + +Carpinus, 346 + Betulus, 66 + +Carthamus, 138 + +Carum, 82, 114, 365 + Bulbocastanum, 159 + Carui, 244, 271, 285, 339, 437 + +CARYOPHYLLACEAE, 99, 113, 120, 137, 139, 140, 148, 250, 379, 397, 398, + 404, 406, 407, 410, 418, 442, 443, 448 + +Casuarina rigida, 325 + +Cassia, 364, 369 + marylandica, 30 + +Castanea vesca, 11, 66, 104, 114, 319, 435 + +Catabrosa aquatica, 351 + +Catalpa, 399 + +Catasetum, 291, 296, 334 + eburneum, 384 + +Cattleya amethystina, 401 + Forbesii, 34, 384, 398 + marginata, 223, 227 + Mossiae, 224, 227, 238 + violacea, 383, 397 + +Caucalis leptophylla, 33 + +Caulophyllum, 75, 125 + +Caylussa, 137 + +Cedrus Libani, 61 + +CELASTRACEAE, 149 + +Celastrus, 149 + +Celosia, 19, 20 + +Centaurea, 37, 114 + collina, 34 + Jacea, 43, 243 + Scabiosa, 20 + moschata, 44 + +Centranthus, 247 + macrosiphon, 250 + ruber, 42, 44 + +Cephalotus follicularis, 314 + +Cerastium, 62, 97, 262, 397, 398 + glomeratum, 280, 358, 463 + tetandrum, 463 + triviale, 252, 280 + vulgatum, 252, 404 + +Cerasus, 74, 117, 149, 250, 260, 424, 489 + avium, 262 + caproniana, 364 + Lauro-cerasus, 64, 66, 370 + vulgaris, 252, 262 + +Ceratonia Siliqua, 30 + +Cercis, 364 + siliquastrum, 325 + +Chamaerops humilis, 300 + +Chaerophyllum, 113 + +Cheiranthus, 121, 131, 136, 148, 295, 364 + Cheiri, 20, 33, 35, 36, 250, 252, 271, 404, 427, 500 + var. gynantherus, 305, 310 + +Cheiranthus incanus, 250 + +Chelone, 361 + barbata, 238 + +Chelidonium majus, 66, 170, 280, 295, 500 + +CHENOPODIACEAE, 397, 405, 406 + +Chenopodium, 45, 365, 367 + glaucum, 397 + murale, 428 + Quinoa, 62, 66 + Vulvaria, 458 + +Chirita sinensis, 170 + +Chlorophytum Sternbergianum, 107 + +Chorozema ilicifolium, 21 + +Chrysanthemum, 16, 72, 365 + indicum, 20, 188, 472 + Leucanthemum, 20 + +Chrysosplenium, 367 + +Cichorium Intybus, 20, 44 + +Cicuta virosa, 406 + +CINCHONACEAE, 429 + +Cionidium Moorei, 190 + +Cirsium, 114, 138 + arvense, 250, 457 + lanceolatum, 20 + tricephalodes, 252, 339 + +Cissus, 211 + +CISTACEAE, 137 + +Cistus vaginatus, 473 + +Citrus, 137, 149, 364, 453 + Aurantium, 33, 35, 44, 56, 75, 134, 303, 310, 335, 369, 388, 389, + 391, 502 + +Clarkia, 24, 295 + elegans, 177, 503 + pulchella, 503 + +Cleistanthus polystachyus, 198 + +Clematis, 136, 148, 288, 295, 367 + florida, 499 + Fortunei, 499 + patens, 499 + Viticella, 28, 499 + +Cleome, 137, 148 + +Cleonia lusitanica, 238 + +Clerodendron fragrans, 506 + +Cliffortia, 396 + +Clinacium dendroides, 49 + +Clitoria Ternatea, 231, 238, 295, 502 + +Clusia rosea, 11 + +Cluytia semperflorens, 198 + +Cneorum, 365 + +Cnicus palustris, 20 + +Cnidium, 113 + +Cobaea scandens, 73, 82, 272, 326, 365 + +Coccoloba platycladon, 328 + +Cochlearia Armoracia, 64, 299, 310, 331 + +Cocos, 115, 365 + nucifera, 429 + +Codiaeum variegatum, 31, 314, 326, 328, 459 + +Coelebogyne ilicifolia, 369 + +Cola acuminata, 370 + +Colchicum autumnale, 45, 67, 73, 250, 287, 296, 406, 407, 507 + +Coleus, 365, 459 + +Columnea Schiedeana, 226 + +Columbine, see _Aquilegia_ + +Colutea, 465 + +Commelyna, 73, 296, 507 + +COMMELYNACEAE, 245, 507 + +COMPOSITAE, 72, 73, 86, 107, 114, 127, 131, 138, 165, 223, 226, 235, + 339, 406, 407, 430, 434, 437, 439, 442 + +Conceveiba macrophylla, 198 + +CONIFERAE, 56, 65, 103, 114, 191, 245, 369, 435 + +Conium maculatum, 114 + +Conostephium, 120 + +Convallaria maialis, 73, 150, 250, 296, 360, 367, 375, 400, 442, 507 + Polygonatum, 507 + +CONVOLVULACEAE, 114, 137, 150 + +Convolvulus, 73, 114, 137, 142, 150, + 296, 510 + arvensis, 20 + Sepium, 20, 108, 250 + tricolor, 505 + +Conyza squarrosa, 20 + +Corallorhiza innata, 238 + +Circeia, 410 + +Corchorus acutangulus, 397 + +Coreopsis, 20, 73, 114, 138 + Drummondi, 339 + +Cornus, 37, 44, 358 + mas, 358, 374 + sanguinea, 44 + suecica, 374 + +Coronilla, 106, 113, 149, 295 + Emerus, 502 + +Correa, 72, 73, 370 + +Cortusa Mathioli, 133, 138, 263 + +Corydalis aurea, 280 + solida, 243 + tuberosa, 235, 336, 238 + +Corylas Avellana, 21, 31, 48, 66, 114, + 349, 354, 364, 365, 368 + +Cotoneaster microphylla, 21 + +Cotula foetida, 19, 20 + +CRASSULACEAE, 113 + +Crassula, 113 + arborescens, 26, 31 + +Crataegus, 82, 113, 149, 295, 364, 404, 419 + Crus galli, 503 + monogyna, 44, 400 + +Crataegus Oxyacantha, 57, 66, 78, 317, 370, 503, 508 + tanacetifolia, 107, 162 + +Crepis, 271 + virens, 20 + +Crinum, 432 + +Crocus, 29, 35, 45, 67, 287, 289, 296, 319, 361, 399, 400, 434, 462 + aureus, 506 + nudiflorus, 302 + pusillus, 506 + vernus, 506 + +Crozophora tinctoria, 198 + +CRUCIFERAE, 73, 76, 98, 113, 136, 139, 141, 148, 257, 297, 364, 379, + 406, 410, 428, 442, 500 + +Cryptomeria japonica, 103, 114, 245, 435 + +Cucubalus, 149 + +Cucumis, 36, 82, 138, 248, 259, 326, 367 + +CUCURBITACEAE, 71, 80, 81, 137, 247 + +Cucurbita, 197, 201, 250, 307, 310, 365, 474 + +Cuphea miniata, 211, 424 + +Cupressus funebris, 218 + +CYCADEAE, 56, 369 + +Cyclamen, 67, 104, 114, 296, 319, 359, 360 + linearifolium, 329 + +Cyclodon, 80 + +Cydonia vulgaris, 71, 79, 295, 423 + japonica, 503 + +Cynanchum fuscatum, 369 + nigrum, 369 + +Cynosurus cristatus, 169 + +CYPERACEAE, 115, 138, 150, 169, 350 + +Cypripedium, 27, 92, 381, 386 + candidum, 401 + Hookerae, 386 + insigne, 91 + +Cyrtanthus, 177 + +Cytisus, 113, 295, 336 + albus, 502 + Laburnum, 21, 66, 157, 189, 222, 226, 231, 238, 356, 459 + nigricans, 15, 104 + + +D. + +Dactylis, 115 + glomerata, 169 + +Dahlia, 44, 72, 188, 244, 433 + variabilis, 20 + +Danae, 60 + +Daphne indica, 21 + odora, 21 + +Datura, 285, 291, 296, 365, 378, 407 + +Datura arborea, 505 + cornigera, 505 + chlorantha, 505 + fastuosa, 296, 379, 450, 505 + humilis, 505 + +Daucus Carota, 53, 57, 82, 113, 121, 124, 125, 137, 149, 244, 250, 252, + 256, 260, 262, 296, 298, 339, 365, 368, 369, 370, 457, 504 + +Delphinium, 44, 136, 148, 225, 283, 286, 295, 344, 364, 367, 374, 388, + 399, 407 + Ajacis, 250, 252, 261, 271, 339, 500 + amoenum, 261 + cheilanthum, 500 + Consolida, 373, 500 + crassicaule, 252, 256, 261, 271, 339 + dictyocarpum, 271, 432 + elatum, 20, 126, 237, 238, 261, 267, 271, 309, 310, 339 + elegans, 500 + grandiflorum, 500 + peregrinum, 219, 226 + +Dendrobium, 227 + nobile, 94, 398 + normale, 224, 383 + +Desmodium canadense, 271 + marylandicum, 467 + +Deutzia, 295 + crenata, 503 + +Dianthus, 67, 113, 121, 129, 137, 139, 145, 146, 149, 166, 261, 268, + 289, 295, 297, 310, 360, 364, 371, 379, 397, 471 + arboreus, 501 + barbatus, 325, 404, 501 + Caryophyllus, 501 + corymbosus, 501 + deltoides, 501 + hybridus, 501 + plumarius, 501 + Poiretianus, 501 + sinensis, 370, 501 + +Dictamnus, 121, 122, 137, 140 + albus, 256, 271 + Fraxinella, 252, 262, 271, 278, 280 + +Dielytra, 236, 237 + +Digitalis lutea, 60 + orientalis, 238 + purpurea, 20, 40, 44, 73, 98, 121, 129, 137, 150, 226, 233, 238, + 296, 298, 315, 365, 373, 398, 459, 472, 505 + +DILLENIACEAE, 398 + +Dionaea, 308 + Muscipula, 310 + +Diosma, 369 + +Diphaca, 365 + +Diplotaxis, 136, 148, 364 + muralis, 252, 458 + tenuifolia, 73, 250, 261, 274, 280, 397, 398, 430 + +DIPSACACEAE, 86, 107, 114, 138 + +Dipsacus, 419, 429 + fullonum, 20, 62, 66, 281, 320, 321, 325 + Gmelini, 325 + pilosus, 20, 325 + sylvestris, 10, 20 + +Dipterocarpus, 249 + +Ditaxis lancifolia, 380 + +Dodecatheon, 138 + +Dodonaea viscosa, 20 + +Draba, 364 + +Dracocephalum austriacum, 238 + moldavicum, 20 + speciosum, 320, 325 + +Dracontium pertusum, 25 + +Drosera intermedia, 170, 265, 271, 277, 473 + + +E. + +Ebenus cretica, 26 + +Eccremocarpus scaber, 222, 226, 326 + +Echeveria, 113 + +Echinophora maritima, 252 + +Echium pyrenaicum, 20 + simplex, 20 + vulgare, 374 + +Echinocactus, 149, 178, 417 + +Elegia, 115 + +Empetrum nigrum, 197 + +Encamptodon perichaetialis, 174 + +EPACRIDACEAE, 120, 137 + +Epacris, 103, 137 + impressa, 61, 379, 504 + +Epidendrum, 114 + elongatum, 107 + Stamfordianum, 401 + +Epimedium, 23, 226 + Musschianum, 390 + +Epiphyllum, 328 + +Epipactis palustris, 325 + +Epilobium, 81, 137, 273 + angustifolium, 20 + hirsutum, 246, 250, 252, 256, 262, + 281, 442 + palustre, 271 + tetragonum, 503 + +Episcia bicolor, 170 + +EQUISETACEAE, 189, 350 + +Equisetum, 61, 325 + fluviatile, 325 + limosum, 325 + Telmateia, 323, 325 + +Eranthis hyemalis, 23, 70 + +ERICACEAE, 114, 119, 137 + +Erica, 21, 73, 137, 296 + cinerea, 504 + hyemalis, 378, 504 + multiflora, 372 + Tetralix, 286, 310, 405, 406, 504 + +Eriobotrya japonica, 295 + +Erodium, 463 + +Ervum Lens, 20, 25, 66 + +Erucago, 136 + +Erucastrum Pollichii, 271 + canariense, 280 + +Eryngium, 113, 368, 442 + viviparum, 104 + +Erysimum, 136, 148 + Barbarea, 252 + cheiranthoides, 252 + officinale, 252 + +Erythrochiton hypophyllanthus, 32, 174 + +Escholtzia crocea, 250 + +Eucomis, 103 + +Eugenia Jambos, 369 + +Euonymus japonicus, 20 + latifolius, 369 + +EUPHORBIACEAE, 114, 150, 369, 395 + +Euphorbia, 114, 150, 365, 371, 395 + Characias, 20 + Cyparissias, 20, 244 + Esula, 198, 307, 310 + exigua, 20 + geniculata, 253, 256 + helioscopia, 56 + Lathyris, 244 + rosea, 369 + palustris, 106, 299 + Peplus, 162 + pusilla, 244 + segetalis, 281 + + +F. + +Faba, see _Vicia_ + vulgaris, 397 + +Fabiana, 237 + +Fagus silvatica, 65, 66, 197, 318, 370, 459 + +Festuca, 115 + nemoralis, 169 + ovina, 169 + +Ficaria, (see _Ranunculus Ficaria_) + ranunculoides, 70, 295, 500 + +FICOIDEAE, 43 + +Ficus Carica, 114, 204, 435 + stipulata, 332 + +Filago, 114 + germanica, 108 + +FILICES, 21, 190, 447 + +Fourcroya, 115 + +Fragaria, xxxvi _adnot._, 250, 295 + alpina, 271 + botryformis, 47 + monophylla, 396 + vesca, 20, 70, 116, 195, 275, 281, 406, 503 + +Fraxinus excelsior, 13, 21, 66, 325, 396, 421 + Ornus, 21 + +Fritillaria imperialis, 21, 45, 296, 462, 506 + Meleagris, 506 + +Fuchsia, 35, 38, 44, 57, 81, 91, 127, 199, 247, 250, 288, 290, 291, + 292, 294, 295, 316, 359, 360, 364, 367, 400, 443 + globosa, 503 +Funckia, 369 + + +G. + +Gagea, 365, 367 + arvensis, 375, 508 + +Gaillardia, 269, 271 + +Galanthus, 296 + nivalis, 300, 506 + +Galeobdolon luteum, 226, 238 + +Galeopsis, 429 + Ladanum, 238 + ochroleuca, 44 + Tetrahit, 429 + +Galium Aparine, 205, 325 + Mollugo, 321, 325 + verum, 325 + +Gaudichaudieae, 334 + +Gardenia, 296, 377 + florida, 504 + Fortuniana, 504 + radicans, 504 + +Gaura biennis, 20 + +Genista, 295 + tinctoria, 502 + sibirica, 502 + Scoparia, 502 + +GENTIANACEAE, 137, 150, 505 + +Gentiana, 71, 73, 137, 150, 252, 296 + Amarella, 166, 305, 310, 371, 505 + campestris, 250, 299, 442 + purpurea, 389 + +GERANIACEAE, 113, 137, 501 + +Geranium, 20, 137, 221, 246, 250, 252, 292, 295, 418 + columbinum, 461 + nodosum, 34, 65 + pratense, 501 + sylvaticum, 501 + +GESNERACEAE, 38, 114, 222, 505 + +Gesnera, 33, 44, 95, 171, 339, 357, 427 + Geroltiana, 88, 89 + zebrina, 355 + +Geum, 121, 137, 465 + coccineum, 275 + rivale, 122, 130, 131, 250, 252, 281, 503 + +Gilia capitata, 281 + glomeruliflora, 253, 271 + +Gladiolus, 21, 296 + tristis, 506 + +Glancium luteum, 66 + +Gleditschia, 30, 177, 364 + triacanthos, 44, 48 + +Glochidion, 310 + +Gloxinia, 171, 206, 207, 222, 226, 238, 284, 291, 296, 365, 451, 506 + +Glyceria aquatica, 169 + fluitans, 169, 278 + +Godetia, 295, 510 + +Godoya, 374 + +Gomphia, 281 + +Gongora, 35 + +Goodenia ovata, 21, 31 + +GRAMINACEAE, 115, 138, 278, 350, 391 + +Gratiola, 296 + +Guarea, 508 + +Gypsophila, 149 + + +H. + +Habenaria, 138, 238 + chlorantha, 382 + +Halenia, 222 + heterantha, 222, 226 + +Hedera Helix, 65 + +Hedypnois, 114 + +Helianthemum, 132, 137, 295, 404 + vulgare, 501 + +Helianthus, 38, 44, 66 + annuus, 20 + tuberosus, 20 + +Helleborus, 23, 288 + foetidus, 442 + olympicus, 284 + +Heliotropium peruvianum, 510 + +Helwingia, 174 + +Hemerocallis, 138, 296, 310, 507 + disticha, 507 + fulva, 507 + +Hepatica, 295, 463 + triloba, 500 + +Heracleum, 82, 113, 137, 150, 262, 365, 368 + Sphondylium, 252, 256, 339 + +Hermesia castaneifolia, 194 + +Herreria parviflora, 141, 150 + +Hesperis, 19, 136, 295 + matronalis, 20, 252, 280, 325, 500 + +Heterocentron, 354 + +Hibiscus, 137, 293, 295, 297, 360, 510 + albus, 501 + flavescens, 501 + Rosa sinensis, 501 + Syriacus, 20, 501 + +Hieracium, 138 + aureum, 17 + praealtum, 339 + Pilosella, 20 + umbellatum, 20 + +HIPPOCASTANEAE, 501 + +Hippeastrum, 296 + equestre, 506 + +Hippuris, 196 + vulgaris, 325, 406 + +Hodgsonia, 326, 474 + +Holeus mollis, 169 + +Honckenya peploides, 196, 406, 461 + +Hordeum, 115, 351 + nepalense, 174, 175 + trifurcatum, 174, 175 + +Humulus Lupulus, 193, 244, 435, 472 + +Hyacinthus, 138, 150, 262, 296, 360, 361, 486 _adnot_ + comosus, 409 + orientalis, 21, 45, 48, 54, 172, 189, 286, 299, 348, 507 + Pouzolzii, 170 + +Hydrangea, 417 + +Hydrocera triflora, 359 + +HYDROCHARIDACEAE, 506 + +Hydrocharis, 296 + morsus ranae, 506 + +Hydrocotyle, 113 + +HYDROPHYLLACEAE, 138 + +Hydrophyllum, 138 + +Hymenocallis, 404 + americana, 462, 463 + +Hyoscyamus, 430 + +Hypericum, 442 + perforatum, 369, 458 + +Hypnum triquetrum, 49 + +Hypochaeris, 73, 138 + radicata, 250, 339, 437, 457 + +Hyssopus officinalis, 20, 325 + + +I. + +Iberis, 295, 364 + amara, 500 + umbellata, 500 + +Ilex Aquifolium, 21, 66, 447 + +Impatiens, 161, 231, 295, 299 + Balsamina, 238, 502 + +Imperatoria, 82 + +Indigofera, 459 + +Inula, 20 + +Ionopsidium acaule, 362 + +Ipomoea, 296, 510 + pandurata, 505 + +IRIDACEAE, 138, 506 + +Irina, 64 + glabra, 65 + +Iris, 138, 286, 296, 359, 360, 361, 365, 401, 430 + Kaempferi, 506 + sibirica, 506 + versicolor, 45 + +Isochilus, 386 + +Ixia carminosa, 84 + +Ixora, 296 + grandiflora, 504 + + +J. + +Jasione, 20, 114 + +JASMINACEAE, 137, 504 + +Jasminum, 137, 296, 360, 400 + grandiflorum, 288, 505 + hirsutum, 505 + nudiflorum, 21 + officinale, 21, 505 + Sambac, 505 + +Jatropha Pohliana, 254, 256 + +Juglans, 244, 400 + nigra, 396 + regia, 66, 193 + +JUNCACEAE, 115, 167, 169 + +Juncus, 115, 317 + articulatus, 431 + conglomeratus, 325 + uliginosus, 107 + +Juniperus virginiana, 194 + sinensis, 217 + +Jussicua, 81, 180 + +Justicia oxyphylla, 25 + + +K. + +Kerria, 295 + +Knautia arvensis, 20, 114 + +Kochia Scoparia, 430 + + +L. + +LABIATAE, 138, 429 + +Laburnum (see _Cylisus_), 65, 157, 189, 222, 226 + +Lacistema, 359 + +Lactuca, 114 + sativa, 11, 20, 33, 44, 313 + +Lambertia, 365 + +Lampsana, 114 + +Lamium, 73, 238, 361 + album, 62, 63, 66, 86, 409 + amplexicaule, 404 + purpureum, 66, 325, 404 + +Larix, 114 + europaea, 21, 90, 245, 435 + microcarpa, 192 + +Lathyrus latifolius, 262 + tuberosus, 30 + +Laurus, 296, 362 + nobilis, 506 + Sassafras, 250, 331, 506 + +Lavatera trimestris, 20 + +Lebeckia, 459 + +Lecythis, 149, 180 + +LEGUMINOSAE, 48, 71, 73, 106, 113, 122, 137, 139, 146, 147, 149, 272, + 276, 429, 434, 444 + +Leitneria floridana, 194 + +Leontice, 125 + +Leontodon, 17, 20, 44, 163, 243, 442 + +Lepidium, 148, 364 + sativum, 57 + +Lepyrodia hermaphrodita, 197 + +Leskea sericea, 49 + +Leucanthemum, 86 + +Leucobryum giganteum, 194 + +Leucoium, 150, 296 + aestivum, 84, 138, 350 + vernum, 350, 506 + +Lilium, 73, 106, 115, 296, 367, 375, 421 + auratum, 73, 289, 400 + bulbiferum, 45 + candidum, 21, 286, 325, 375, 507 + cruentum, 21 + lancifolium, 35, 400, 443 + longiflorum, 310 + Martagon, 21, 286, 325, 507 + tigrinum, 306, 310 + +Linaria, 137, 229, 230, 233, 296, 316, 361, 365, 367, 405 + aeruginea, 238 + chalepensis, 238 + Cymbalaria, 238 + decumbens, 238 + Elatine, 238 + origanifolia, 238 + Pelisseriana, 238 + pilosa, 238 + purpurea, 20, 44, 238 + spuria, 238 + triphylla, 238 + vulgaris, 162, 226, 234, 235, 238, 316, 374, 505 + triornithophora, 238 + +Linum, 335 + +Linum usitatissimum, 20 + +Liquidambar, 362 + +Listera ovata, 398 + +LOBELIACEAE, 72, 114 + +Lobelia, 211, 424 + +Lolium, 86, 113, 115, 351 + perenne, 61, 169, 325 + +Lonicera, 38, 44, 73, 82, 226, 271, 281, 296, 297 + brachypoda, 358 + Caprifolium, 408 + Periclymenum, 66, 226, 251, 256, 262, 263, 338, 379, 404, 406, 504 + Xylosteum, 252, 358 + +Lopezia, 298, 410 + +Lotus, 113, 295, 360, 510 + corniculatus, 104, 377, 436, 502 + uliginosus, 363 + +Lowea, 396 + +Lunaria, 136, 364 + +Lupinus, 106, 165, 226, 280 + polyphyllus, 238 + +Lycaste Skinneri, 93, 95, 291 + +Lychnis, 113, 137, 148, 295 + chalcedonica, 501 + Coronaria, 107, 510 + dioica, 67, 252, 262, 280, 404, 464 + flos cuculi, 501 + sylvestris, 252, 501 + vespertina, 501 + Viscaria, 501 + +Lycium, 365, 367 + europaeum, 250 + +Lycopersicum, see _Solanum_ + esculentum, 389 + +Lysimachia, 119 + Ephemerum, 271, 281 + nummularia, 505 + vulgaris, 20, 87 + +Lythrum, 335 + Salicaria, 374 + + +M. + +Maelenia, 403 + +Maesa, 145, 371 + +MAGNOLIACEAE, 122 + +Magnolia, 288, 388, 440 + Campbelli, 427 + fuscata, 304, 310, 427 + +MALPIGHIACEAE, 137, 334, 403, 404, 406 + +Malus, 78, 79, 388, 389, see _Pyrus_ + +MALVACEAE, 137, 149, 288, 292, 295, 362, 395 + +Malva, 295 + crispa, 448 + moschata, 501 + rotundifolia, 501 + sylvestris, 252 + +Mangifera, 159, 369 + +Marchantia, 174 + +Marcgraavia, 23 + umbellata, 332 + +Masdevallia, 27, 95 + +Mathiola, 136, 148, 295 + annua, 361, 500 + incana, 20, 38, 44, 68, 299, 500 + glabrata, 500 + +Matricaria Parthenium, 281 + +Maxillaria, 383 + +Medicago, 137, 149, 262, 295, 364, 502 + lupulina, 404, 432 + maculata, 218, 271 + +Melastoma, 74, 125, 424 + +Melia Azedairach, 21 + +Melianthus major, 298, 407 + +Melilotus, 137, 139, 147, 149, 262 + arvensis, 374 + leucantha, 432 + macrorhiza, 271 + officinalis, 404 + +Melittis, 361 + +Mentha, 238, 361, 459 + aquatica, 325 + viridis, 325 + +Mercurialis, 62, 194, 365 + annua, 198 + perennis, 66 + +Mesembryanthemum, 26 + +Metrosideros, 103 + +Miconia, 355 + +Mimosa, 365 + Lophantha, 31 + +Mimulus, 73, 284, 296 + luteus, 505 + +Mnium serratum, 49 + +Mirabilis, 296, 418 + Jalapa, 506 + +Modecca, 326 + +Mollugo Cerviana, 398 + +Momordica Elaterium, 20 + +Monarda fistulosa, 298 + +Morus, 193, 354, 459 + +Mozinna, 399 + peltata, 194 + +Musa, 245, 407 + +Muscari, 408, 461 + comosum, 340, 467 + +MUSCI, 174 + +Mussaenda, 249, 283, 429 + +Myosotis caespitosa, 281 + palustris, 375 + scorpioides, 20 + +Myosurus, 440 + +Myristica moschata, 194 + +MYRTACEAE, 80, 149, 362, 395 + +Myrtus, 295, 310 + communis, 503 + + +N. + +Narcissus, 21, 67, 138, 286, 296, 360, 365, 443 + aureus, 506 + biflorus, 34, 45, 506 + chrysanthus, 38, 45 + concolor, 506 + Cypri, 506 + Jonquilla, 506 + incomparabilis, 38, 45, 301, 506 + italicus, 506 + lobularis, 506 + major, 154 + montanus, 301 + poculiformis, 506 + poeticus, 24, 33, 301, 506 + pseudo-narcissus, 506 + Tazetta, 45, 300, 506 + Telamonius, 506 + +Nasturtium, 136, 271 + amphibium, 181 + +Nelumbium, 295 + speciosum, 500 + +Nepeta diffusa, 238 + +Nepenthes, 328, 473 + +Nephrodium molle, 447 + +Nerium Oleander, 62, 66, 296, 301, 377 + odorum, 505 + +Neuropeltis, 32 + +Nicandra physaloides, 458 + +Nicotiana, 23, 31, 73 + rustica, 281 + +Nigella, 220, 262, 295, 297, 364, 374, 399 + damascena, 226, 269, 271, 286, 375, 500 + +Nuytsia, 371 + +NYMPHAEACEAE, 148 + +Nymphaea, 50, 141, 143, 148, 285, 295 + alba, 162 + dentata, 256, 261 + guineensis, 170 + Lotus, 277, 280 + + +O. + +Octadenia, 364 + +Odontoglossum Alexandrae, 387, 403 + Uro Skinneri, 463 + +OEnanthe, 82, 114, 360, 365, 457 + crocata, 80, 359, 457 + +OEnothera, 44, 367 + striata, 252 + +Olea europoea, 21, 157, 420 + +ONAGRACEAE, 81, 137, 406, 503 + +Oncidium, 67, 114, 227 + abortivum, 462 + bicolor, 45 + Cebolleta, 107 + +Oncidium cucullatum, 91 + heteranthum, 224 + ornithorhyncum, 43 + +Ononis minutissima, 404 + monophylla, 396 + +Opercularia, 38 + +Ophrys, 114 + apifera, 360, 398 + aranifera, 35, 42, 45, 238, 298, 383, 384, 385, 386, 398 + insectifera, 27, 301, 382 + fucifera, 506 + +Opuntia, 81, 113, 149, 180 + fragilis, 107, 178 + monacantha, 178 + Salmiana, 107, 178, 179 + +ORCHIDACEAE, 27, 34, 42, 91, 112, 114, 128, 138, 150, 153, 209, 223, + 290, 345, 360, 366, 367, 380-387, 397 + +Orchis, 138, 150, 153, 154, 227, 296 + conopsea, 238 + latifolia, 238 + mascula, 153, 154, 238, 299, 387, 506 + militaris, 387 + Morio, 238, 382, 384, 387, 506 + palustris, 366 + papilionacea, 238 + pyramidalis, 128, 238, 506 + sambucina, 63, 66 + simia, 238 + +Ornithogalum, 114, 360, 365, 367 + longebracteatum, 171 + +OROBANCHACEAE, 137 + +Orobanche, 73, 137, 209, 296, 505 + gracilis, 442 + +Orobus, 295 + vernus, 502 + viscosus, 502 + +Ouvirandra, 458 + +OXALIDACEAE, 403 + +Oxalis, 295, 329, 404, 459 + Acetosella, 404 + carnea, 502 + + +P. + +Paeonia, 295, 300, 364, 399, 407 + Moutan, 500 + officinalis, 261 + paradoxa, 500 + tenuiflora, 500 + +PALMACEAE, 155 + +Panax, 331 + +Papaver, 289, 295, 297, 473 + bracteatum, 28, 304, 310, 500 + nudicaule, 310 + orientale, 250 + +Papaver Rhoeas, 429, 458, 500 + somniferum, 28, 305, 310, 500 + +PAPAVERACEAE, 99, 500 + +PAPILIONACEAE, 397, 502, see _Leguminosae_ + +Paris quadrifolia, 31, 358, 360, 367, 396 + +Paritium, 137 + +Parnassia, 364 + +Paronychia serpyllifolia, 473 + +Parthenium inodorum, 243 + +Passiflora, 137, 201, 301, 365, 463, 473 + coerulea, 185 + palmata, 185 + quadrangularis, 181 + +PASSIFLORACEAE, 137 + +Pastinaca, 82 + sativa, 272 + +Pavia, 137 + +Pedicularis, 238 + euphrasioides, 238 + sylvatica, 44, 223, 226, 238 + +Peganum, 363 + +Pelargonium, 22, 23, 25, 63, 107, 113, 208, 221, 225, 295, 336, 360, + 373, 389, 418, 419, 434 + grandiflorum, 86 + inquinans, 30, 226 + zonale, 106, 226, 501 + +Peltaria, 137 + alliacea, 250, 251, 252 + +Pentstemon, 226, 455 + +Pereskia, 107, 113, 149 + Bleo, 179 + +Perilla, 459 + +Persica (_Amygdalus_) + vulgaris, 44, 250, 252, 503 + +Petalostylis, 296 + +Petunia, 254, 256, 291, 296 + nyctaginiflora, 505 + violacea, 34, 44, 250, 253, 505 + +Phaius grandiflorus, 107, 509, 510 + +Phalaenopsis, 238 + amabilis, 227 + equestris, 231, 238 + Schilleriana, 224 + +Pharbitis, 73 + +Phaseolus, 21, 298, 364 + +PHILADELPHACEAE, 113, 137, 149 + +Philadelphus, 113, 137, 142, 149, 295, 364, 367 + coronarius, 503 + speciosus, 178, 298 + +Philyra brasiliensis, 198 + +Phleum, 115, 138 + phalaroides, 169 + pratense, 325 + +Phlomis, 138 + biloba, 72 + fruticosa, 119, 238 + +Phlox, 73 + +Phoenix dactylifera, 318 + +Phylica, 325 + +Phyllanthus longifolius, 198 + +Phyllarthron, 64, 396 + +Phyteuma odoratum, 271 + orbiculare, 20 + spicatum, 166, 252 + +Phytolacca, 20 + +Picea, see _Abies_ + Lowii, 203 + nobilis, 203 + Nordmanniana, 203 + Webbiana, 203 + +Picrasma, 509 + +Pimpinella magna, 66 + Saxifragra, 66 + +Pinckneya, 429 + +Pinus, 11, 19, 114, 318, 346, 471, (see _Abies_, _Larix_, _Cedrus_) + alba, 192 + pinea, 90, 218 + Pinaster, 13, 21 + sylvestris, 21, 349 + +Pistacia Lentiscus, 197 + +Pisum, 137, 295 + sativum, 27, 30, 31, 95, 166, 302, 372, 432, 472, 502, 510 + +Platycodon, 296 + grandiflorum, 504 + +PLANTAGINACEAE, 114 + +Plantago, 114, 136 + Coronopus, 458 + lanceolata, 61, 104, 108, 110, 111, 243 + media, 20, 111 + major, 65, 108, 109, 111, 243, 372, 374 + maritima, 108 + +Platanus, 50 + +Plectranthus fruticosus, 238 + +PLUMBAGINACEAE, 114 + +Plumbago, 360 + +Poa alpina, 169 + annua, 169 + bulbosa, 169 + pratensis, 169 + trivialis, 169 + +Podalyria myrtillifolia, 44 + +Podocarpus, 426 + +Podophyllum, 295 + peltatum, 500 + +Podospermum laciniatum, 247, 250 + +Poggendorffia, 294 + +Pogonia ophioglossoides, 238, 386 + +Poinsettia, 340 + +Polemonium coeruleum, 20, 66, 253, 305, 310, 404 + +Polianthes, 296 + +Polygala vulgaris, 399 + +POLYGONACEAE, 114, 138, 150 + +Polygonatum anceps, 42, 45 + multifolium, 23, 31 + +Polygonum, 114, 365 + orientale, 31, 390 + viviparum, 106, 169 + +Polypodium anomalum, 190 + +POMACEAE, 70, 71, 77, 79, 142, 405, 503 + +Pomaderris elliptica, 355 + +Pomax, 38 + +Populus, 202, 309 + alba, 66 + +Portulaca, 295 + +Potamogeton, 329 + bifolium, 434 + +Potentilla, 139, 140, 149, 295, 374, 438 + alpestris, 503 + argentea, 280 + anserina, 503 + major, 374 + nepalensis, 250, 252, 275, 280 + reptans, 503 + Tormentilla, 503 + +Poterium, 113 + Sanguisorba, 107 + polygamum, 281 + +Primula, 71, 73, 114, 121, 138, 150, 296, 335, 360, 365, 367 + acaulis (see _vulgaris_), 45, 248, 250, 308, 310, 377, 504 + Auricula, 45, 133, 271, 282, 504 + denticulata, 20, 504 + elatior, 45, 250, 504 + calycanthema, 283 + imperialis, 103 + officinalis, 250, 283 + praenitens, 271, 504 (see _sinensis_) + sinensis, 24, 45, 95, 98, 103, 133, 253, 256, 262, 263, 271, 281, + 297, 314, 315, 389, 429, 441, 449 + variabilis, 104 + veris, 17, 20, 35, 39, 45, 105, 164 + villosa, 504 + vulgaris, 17, 70, 106 (see _acaulis_ and _officinalis_) + +PRIMULACEAE, 98, 114, 118, 121, 133, 138, 150, 262, 339, 366, 504 + +Prismatocarpus, 114, 150, 178 + +Prunella vulgaris, 226 + +Prunus, 137, 203, 295, 300, 360, 364, 367 (see _Cerasus_, _Amygdalus_, + _Persica_) + Armeniaca, 44 + Cerasus, 74 + domestica, 366, 464 + Lauro Cerasus, 21, 446 + spinosa, 44 + +Prunus sylvestris, 21 + +Pseudostachyum polymorphum, 168 + +Psittacanthus, 371 + +Ptelea, 364 + +Pterandra, 290 + +Pteridophyllum, 331 + +Pterisanthes, 328 + +Pteris quadriaurita, 333 + +Punica Granatum, 21, 295, 325 + +Pyrethrum, 114 + inodorum, 431 + Parthenium, 339 + +Pyrus, 82, 107, 113, 121, 137, 149, 203 + communis (Pear), 36, 66, 162, 178, 422, 423, 503 + dioica, 304 + japonica, 161 + Malus (Apple), 44, 78, 79, 166, 210, 250, 282, 295, 310, 325, 327, + 375, 406, 420, 503 + spectabilis, 508 + torminalis, 325 + + +Q. + +Quercus, 51 + Cerris, 66 + pubescens, 66 + + +R. + +Raphanus sativus, 161, 252, 327, 360, 369 + +RANUNCULACEAE, 113, 122, 136, 148, 195, 246, 339, 410, 499 + +Ranunculus, 19, 55, 113, 119, 258, 288, 295, 296, 297, 360, 367, 407 + aconitifolius, 500 + acris, 67, 250, 436, 500 + aquatilis, 458 + asiaticas, 500 + auricomus, 195, 285, 397, 406, 461, 469 + bulbosus, 17, 20, 44, 195, 406, 500 + bullatus, 500 + Ficaria, 70, 96, 195, 261, 368, 406 + fluitans, 433, 437 + gramineus, 500 + Lingua, 44, 67 + Philonotis, 20, 339 + tripartitus, 20, 46 + repens, 23, 252, 261, 500 + +RESEDACEAE, 137, 148 + +Reseda, 67, 137, 141, 148 + lutea, 252, 271 + Luteola, 61, 112, 399 + odorata, 20, 44, 280 + Phyteuma, 252, 261 + +RESTIACEAE, 115, 167, 189, 350, 463 + +Restio, 115, 194 + +Rhamnus catharticus, 463 + Frangula, 280 + +Rhinanthus crista galli, 238 + +Rhodiola rosea, 197 + +Rhodora, 73 + +Rhododendron, 72, 73, 91, 133, 137, 176, 289, 290, 296, 453 + indicum, 504 + linearilobum, 72 + +Rhus Cotinus, 62, 66, 348, 409, 467, 472 + Toxicodendron, 66 + +Ribes, 296 + nigrum, 66, 296 + sanguineum, 503 + +Richardia aethiopica, 337, 358 + +Ricinus communis, 198, 300 + +Ricotiana, 364 + +Robinia, 365 + Pseudacacia, 44, 204, 317, 325, 396 + +ROSACEAE, 70, 71, 77, 80, 113, 121, 122, 137, 149, 246, 274, 405, 406, + 502 + +Rosa, 21, 77, 79, 82, 113, 121, 130, 137, 139, 149, 151, 152, 162, + 176, 184, 185, 186, 204, 246, 247, 250, 257, 258, 262, 271, 274, + 288, 289, 295, 367, 368, 394, 473, 502 + alpina, 209 + arvensis, 307, 310, 404 + Banksiae, 502 + berberifolia, 396 + canina, 502 + centifolia, 31, 502 + Carolina, 502 + cinnamomea, 502 + damascena, 502 + diversifolia, 280 + Eglanteria, 502 + gallica, 31, 502 + indica, 502 + moschata, 502 + nivea, 502 + pimpinellifolia, 502 + rubiginosa, 502 + spinosissima, 502 + sulphurea, 502 + +RUBIACEAE, 45, 80 + +Rubia tinctorum, 322, 325 + +Rubus, 137, 252, 279, 399, 406, 429, 465 + arcticus, 67 + caesius, 281, 285, 502 + corylifolius, 502 + fruticosus, 66, 281, 374, 380, 461, 502 + Idaeus, 396 + rosifolius, 502 + +Rudbeckia, 82, 114, 127 + +Ruellia clandestina, 403, 404 + +Rumex, 45, 138, 150, 325 + arifolius, 278, 281 + crispus, 304, 310, 448 + scutatus, 278, 281, 431 + +Ruscus, 328, 470 + aculeatus, 60, 318, 348, 445 + +Russellia juncea, 20 + +RUTACEAE, 137, 149 + +Ruta, 363, 367, 371 + + +S. + +Sagina, 397 + procumbens, 501 + +Sagittaria, 296, 329 + latifolia, 507 + sagittifolia, 325, 507 + +Salisburia adiantifolia, 61 + +Salix, 166, 168, 244, 310, 326, 349 + babylonica, 202, 257, 262, 299, 309, 326 + calyculata, 29 + capraea, 271, 299 + cinerea, 41, 45, 189, 299 + fragilis, 357 + monandra, 29 + nigricans, 299 + pendula, 357 + repens, 194 + silesiaca, 299 + vitellina, 21 + +Salpiglossis straminea, 44 + +Salvia, 226 + officinalis, 66 + pratensis, 36 + Verbenaca, 87 + +Sambucus, 296, 359, 360, 365, 367 + nigra, 21, 55, 66, 164, 325, 396 + racemosa, 66 + +Sanguinaria, 295 + canadensis, 500 + +Sanguisorba, 113 + officinalis, 107 + +SANTALACEAE, 122, 138, 150 + +SAPINDACEAE, 137, 448 + +Saponaria, 28, 67, 76, 149, 295, 297, 300, 360, 379 + officinalis, 97, 301, 501 + +Sarothamnus, 360, 363 + Scoparius, 363 + +Saxifraga, 33, 44, 106, 113, 296, 390 + crassifolia, 307, 310 + decipiens, 288 + foliosa, 281 + granulata, 298, 504 + irrigua, 20 + longifolia, 404 + +Saxifraga mutata, 20 + +SAXIFRAGACEAE, 80, 81 + +Scabiosa, 35, 106, 114, 138, 141, 296, 297, 429 + agrestis, 252 + arvensis, 325 + atropurpurea, 62, 66 + Columbaria, 252, 281 + +Scandix, 125 + +Schismatopera distichophylla, 194 + +Schoenodorus, 364 _adnot_ + +Schoenus cephalotes, 245 + +Schoenodon, 365 + +Sciadopitys verticillata, 352 + +Scilla, 296 + autumnalis, 507 + nutans, 507 + +Scirpus lacustris, 325 + +Scolopendrium D'Urvillei, 332 + vulgare, 64, 314, 326, 328, 459, 467 + +Scorzonera, 19 + octangularis, 247, 250 + +Scrophularia, 114, 455, 463 + aquatica, 226, 238, 262, 281 + arguta, 334 + nodosa, 44, 106, 281 + +SCROPHULARIACEAE, 114, 121, 137, 150, 222, 505 + +Secale cereale, 55, 115 + +Sechium edule, 81, 202 + +Sedum cristatum, 18, 20 + reflexum, 20 + +Selenipedium caudatum, 224, 227 + +Selinum, 82, 114 + caruifolium, 281 + +Sempervivum, 105, 113, 204, 509 + montanum, 310 + sediforme, 164 + tectorum, 292 _adnot_, 308, 310 + +Senecio, 44, 114 + vulgaris, 247, 250, 252, 339, 432, 437, 439, 457 + +Serissa, 296, 377, 504 + +Sesamum, 365 + indicum, 238 + +Seseli, 113, 281 + coloratum, 166 + +Sideritis, 238, 365 + +Silaus, 113, 365 + +Silene, 44, 137, 148, 295, 301 + conica, 68 + Otites, 464 + inflata, 501 + +Sinapis, 57, 136, 270, 295, 371 + arvensis, 181, 250, 261, 264, 271, 280, 501 + +Sinningia purpurea, 431 + +Sisymbrium, 136 + officinale, 250, 261, 271, 280 + +Sisymbrium tenuifolium, 252 + +Slateria, 74 + +SOLANACEAE, 150, 430, 505 + +Solanum, 73, 125, 150, 296, 360, 365, 367, 370, 424, 453 + amazonicum, 430 + Dulcamara, 66, 288, 430, 504 + esculentum, 74 + cornutum, 430 + Lycopersicum, 38, 44, 74, 171, 391, 442 + tridynamum, 430 + tuberosum, 54, 142, 288, 333, 420 + Vespertilio, 430 + +Sophora, 55 + +Spartianthus, 295 + junceus, 502 + +Spartium, 295 + junceum, 18, 21 + Scoparium, 21 + +Spathiphyllum, 245 + +Spilanthes, 138, 365 + oleracea, 44 + +Spinacia oleracea, 31, 197 + +Spiraea, 21, 137, 295 + Filipendula, 503 + oblongifolia, 252 + prunifolia, 503 + Reevesii, 503 + strobilacea, 503 + Ulmaria, 503 + +Spiranthes, 319 + +Splachnum vasculosum, 49 + +Stachys, 138, 339, _adnot_ + germanica, 310 + lanata, 375 + sylvatica, 74, 253, 271, 281 + +Stackhousia juncea, 334 + +Stangeria paradoxa, 172 + +Stapelia, 20 + +Staphylea pinnata, 30 + +Stauntonia latifolia, 194 + +Stellaria, 148, 404, 406 + media, 271, 279, 280 + +STELLATAE, 396, 408 + +STERCULIACEAE, 362, 464 + +Sterculia platanifolia, 20, 256 + +Sternbergia, 296 + lutea, 506 + +Stratiotes aloides, 406 + +Strelitzia juncea, 459 + regina, 25 + +Streptocarpus Rexii, 15, 44, 226, 227, 461 + +Strophanthus, 326 + +Suaeda, 365 + fruticosa, 430 + maritima, 20, 21 + +Suregada, 198 + +Symphoricarpus racemosus, 66 + +Symphyomyrtus, 38 + +Symphytum, 71, 138, 365 + officinale, 253, 262, 263, 271 + Zeyheri, 271 + +Syringa, 296, 360, 367 + persica, 44, 61, 66, 284, 505 + vulgaris, 79, 505 + + +T. + +Tabernaemontana, 296 + coronaria, 504 + +Tacsonia pinnatistipula, 294 + +Tamus communis, 21 + +Taraxacum, 164 (see _Leontodon_) + +Taxodium, 114 + distichum, 444 + +Taxus baccata, 11, 21, 90 + +Tetragonia, 113 + expansa, 142, 180 + +TETRAGONIACEAE, 149 + +Teucrium campanulatum, 226, 233, 238 + +Thalictrum, 205, 500 + minus, 307, 310 + +Thea, 295, 502 + +Thelymitra, 224 + +Thesium, 121, 123, 138, 150 + +Thlaspi arvense, 250, 271 + bursa-pastoris, 252, 428 + +Thuja occidentalis, 319 + orientalis, 21 + +Thymus Serpyllum, 325, 405, 406 + +Thysselinum, 137, 365 + palustre, 437, 457 + +Tigridia, 360, 361, 365 + Pavonia, 35, 462 + +Tilia asplenifolia, 66, 459 + europaea, 22, 30, 65 + parvifolia, 66 + +Tiliaceae, 137, 362 + +Tithonia, 26 + +Tofieldia, 296 + calyculata, 300 + +Torenia scabra, 406 + +Torilis, 82, 149 + Anthriscus, 256, 271, 281, 339, 406, 441, 442 + +Tradescantia, 245, 296, 360, 365, 367, 404 + virginica, 88, 507 + +Tragopogon, 19, 20, 73, 114, 431 + orientale, 250 + pratense, 247, 250, 442, 457 + +Trapa natans, 202, 364 + +Trichostomum rigidulum, 49 + +Trifolium, 106, 113, 121, 137, 139, 149, 295, 360, 364 + +Trifolium hybridum, 262, 399, 406, 407 + pratense, 20 + repens, 20, 23, 68, 70, 98, 145, 146, 226, 231, 238, 250, 252, 256, + 260, 262, 265, 271, 276, 279, 356, 368, 397, 399, 406, 407, 434, + 436, 438, 502 + resupinatum, 20 + +Trillium grandiflorum, 507, 510 + +Trinia vulgaris, 405 + +Triphasia aurantiaca, 369 + +Triticum, 86, 115, 350, 391 + repens, 106, 325 + vulgare, 55, 113 + +Triumfetta, 137, 252, 259, 260, 262, 265, 271, 280 + +Trollius europaeus, 66, 295, 500 + +TROPAEOLACEAE, 149, 501 + +Tropaeolum, 149, 398, 283 + majus, 20, 222, 225, 226, 232, 238, 251, 252, 254, 256, 271, 280, + 295, 310, 406, 442, 501 + minus, 501 + +Tulipa, 35, 45, 67, 75, 84, 85, 138, 262, 300, 302, 348, 359, 360, + 361, 365, 367, 388, 390, 421 + Gesneriana, 31, 250, 310, 315, 442, 507 + sylvestris, 507 + +Turritis, 271 + glabra, 252 + +Typha, 189 + + +U. + +Ulex, 295, 360 + europaeus, 377, 502 + +Ulmus americana, 66 + campestris, 31, 52, 62, 157, 325, 353, 427 + +UMBELLIFERAE, 37, 71, 73, 80, 107, 113, 121, 127, 131, 132, 137, 139, + 140, 143, 149, 150, 162, 244, 257, 261, 339, 358, 395, 405, 406, + 407, 437, 439 + +Uredo candida, 279 + +URTICACEAE, 114 + +Urtica dioica, 62, 66, 194 + +Usteria, 283 + + +V. + +Vaccinium, 68 + +Valantia cruciata, 44 + +VALERIANACEAE, 114 + +Valeriana, 114, 165, 419 + dioica, 325 + officinalis, 325 + +Valisneria spiralis, 319, 433 + +Verbascum, 73, 116, 137, 225, 253, 430 + australe, 35 + nigrum, 226, 251 + phlomoides, 281 + Thapsus, 404 + +Verbena, 67, 68, 506 + +Veronica, 60, 150, 296, 334, 360, 361, 365, 375, 505 + austriaca, 66 + Beccabunga, 468 + Chamaedrys, 442 + latifolia, 459 + longifolia, 325 + spicata, 325, 374 + +Viburnum, 44, 296, 417 + Opulus, 473 + +Vicia, 30, 365, 369, 472 + Faba, 434 + +Vinca, 137, 296 + herbacea, 389 + major, 505 + minor, 44, 99, 358, 390, 505 + rosea, 31 + +Viola, 23, 137, 225, 229, 289, 295, 297, 405 + grandiflora, 501 + hirta, 226, 238 + odorata, 20, 94, 220, 226, 238, 286, 404, 429, 501, 508 + tricolor, 461, 501 + +VIOLACEAE, 137, 334, 403, 406, 500 + +Viscum album, 56, 369, 509 + +VITACEAE, 137 + +Vitex incisa, 238 + +Vitis, 417 + vinifera, 20, 29, 34, 44, 66, 137, 157, 182, 183, 211, 280, 374, + 422, 424 + + +W. + +Wedelia perfoliata, 442 + +Weigela rosea, 358 + +Welwitschia, 162, _adnot_ + +Wildenovia, 115, 167, 168 + +Wistaria, 364 + sinensis, 226 + + +X. + +Xanthosoma appendiculatum, 31 + +Xanthoxylum, 21 + +Xylophylla, 328 + + +Y. + +Yucca, 361 + flexilis, 361 + + +Z. + +Zamia, 170 + +Zea Mays, 21, 113, 136, 191, 197, 300, 310, 350, 369, 466 + +Zingiber Zerumbet, 224, 227 + +Zinnia, 44 + elegans, 20 + +Zygopetalum maxillare, 398 + + + + +ERRATA. + + +The reader is requested to make the following corrections:-- + +Page 182. Fig. 94 should be 94*. + +Page 194. The reference 3 applies not to the nutmeg but to the hop, +figured at p. 193. + +Page 309. Fig. 165 legend--for _Sempervivun tecotorum_ read _Sempervivum +tectorum_. + + + +PRINTED BY J. E. ADLARD, BARTHOLOMEW CLOSE. + + + + +Transcriber's Notes: + +Page xx and 202: +208. Passage of pinnate to palmate leaves in horse-chesnut 439 +'chesnut may be old spelling for chestnut?'. Changed as most are +spelled chestnut. + +Page 65: +Naturforscherversammlung +Changed Naturvorschefversamlung to Naturforscherversammlung to match +other occurrence. (See footnote 528). + +Page 145: +So far as the andraecium is concerned, the stamens +Changed to androecium to match other occurrences. + +Page 149: +Echinocactus changed from Echinocatus to match other +occurrences. + +Page 397: +The species mentioned are _Ambrina ambrosiodes_, +Changed to ambrosioides to match index page. + +Page 502: +Medicago sp., ? ., Europe. +The ? mark replaces a blank in original for a missing attribution. + +Page 503: +ONAGRARIAE changed to ONAGRACEAE to match other occurrences, especially +the index referrence. + +Footnote 126: +'Neue Denkschriften der allgemeine Schweizerischen +Gesellschaft,' Perhaps this should be: allgemeinen. Unchanged. + +Index +Alstroemeria, 319, 326 +Changed to Alstroemeria to match referenced pages. + +DIPSACACEAE, 86, 107, 114, 138 +All dipsaceae changed to dipsacaceae to match index and current spelling. + +Errata changes listed were made. + +Inconsistent hyphenation: +co-existent and coexistent +Lauro-cerasus and Laurocerasus +mid-rib and midrib +outgrowth and out-growth + + + +***END OF THE PROJECT GUTENBERG EBOOK VEGETABLE TERATOLOGY*** + + +******* This file should be named 23354.txt or 23354.zip ******* + + +This and all associated files of various formats will be found in: +https://www.gutenberg.org/dirs/2/3/3/5/23354 + + + +Updated editions will replace the previous one--the old editions +will be renamed. + +Creating the works from public domain print editions means that no +one owns a United States copyright in these works, so the Foundation +(and you!) can copy and distribute it in the United States without +permission and without paying copyright royalties. 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