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VAriM «? 



VENDU EN 1022 




This volume, is issued to the Subscribers to the Ray Society/™ 
the Year 1868. 






















D.C.L., LL.D., F.R.3., F.L.S., ETC. ETC. 










PART I. — Union of Organs . . . .8 

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. 

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, 38 — Synanthy, 36 — Syn- 
carpy, 45 — Synspermy, 50 — Between axes of different plants 
of same species, 50 — And of different species. 55 — Synophty, 56. 



PART II. — Independence of Organs . .58 


Fission . . . . . . .59 

Fission of axile organs, 60 — Of foliar organs, 61 — Of petals, 
66 — Of stamens, 68 — Of carpels, 68. 


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. 


Solution . . . . . .76 

Solution of calyx from ovary, 77 — Of stamens from petals, 

PART III. — Alterations of Position . 83 


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. 


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. 




Heterotaxy ...... 150 

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 gemma) 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 Conipositse, 188 — Heterotaxy affecting the inflorescence, 188 
— Supra- soriferous ferns, 190. 


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. 

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. 



PART I. — Stasimorphy . . . .216 


Persistence of juvenile forms .... 217 

Stasimorphy in leaves of Conifers, 217— Regular peloria, 219. 



PART II. — Pleiomorphy . . . 228 


Irregular peloria ..... 228 

PART III. — Metamorphy . . . .240 

Phyllody ...... 241 

Phyllody of bracts, 242 — In inflorescence of Conifers, 245 — 
Of calyx, 245— Of coroUa, 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. 


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 

Deformities ...... 311 

Formation of tubes, 312— In flower, 314— Spurs, 315— Con- 
tortion, 316 — Spiral torsion, 319 — Of leaf, 326 — Adventitious 
tendrils, 326 — Interrupted growth, 327 — Cornute leaves, 328 — 
Flattening, 328. 



POLYMORPHY ...... 329 

Heterophylly. 330 — Dimorphism, 333 — Spoils or bud-varia- 
tions, 336. 


Alterations of colour ..... 337 

Albinism, 337 — Virescence, 338— Chromatism, 339. 



PART I.— Increased Number ok Organs . 343 

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. 

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 andrcecium, 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 — Andrcecimn, 379 — Androecium 
of Orchids. 380— Pleiotaxy of gyncecium, 388— Increased num- 
ber of flowers in an inflorescence. 391. 



PART II. — Diminished Number of Organs . 392 


Suppression of axile organs .... 393 

Acaulosia, 393 — Non- development of peduncle, 393 — Nature 
of calyx-tube (Casimir de Candolle), 394 — Suppression of 
columella, 395. 


Suppression of foliar organs .... 395 

Aplrylly, 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 andrceciurn, 405 — Of gynoecium, 406 — Of ovules or seeds, 407 
Of flower, 408 — General remarks on suppression, 409. 


ENCE . . . . .413 

PART I. — Hypertrophy . 416 


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 — En- 
largement of perianth, 428 — Of andrceciurn, 430— Of gynoecium, 
430 — Of fruit, 431 —Alterations of consistence, 432. 


Elongation ...... 433 

Elongation of root, 434 — Of inflorescence, 434 — Of flower- 
stalks, 435— Of leaves, 437— Of parts of flower, 438— Of thala- 
mus and placenta, 440 — Apostasis, 440. 

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, $■'>'■'>. 

PART II.— Atrophy . . . lw 


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— De- 
pauperated ferns, 466 — General remarks, 467. 

Degeneration ...... 470 

Formation of scales, 470 — Of hairs, 472 — Of glands, 473 — Of 
tendrils, 473. 


General morphology of the leaf and axis, 476. Homology. 
476— Special morphology, 479— Calyx-tube, 480— Androecinm, 
482— Inferior ovaiy. 482— Plaeentation, 483— Structure of the 



ovule, 484 — Leaves of Conifers, 484 — Relative position of 
organs, 484 — Law of alternation, 485 — Corelation, 486 — Com- 
pensation, 488 — Teratology and classification, 488. 


Double flowers, varieties of, 491 — Causes of production, 491— 
Relation to vai-iegated foliage, 497 — List of plants producing, 

NOTE . . . . . .508 













Diagram of regular pentamerous flower . . .4 

Cohesion of branch Dipsacus sylvestris . . .10 

Fasciated lettuce (' Gard. Chron.') . . .11 

Fasciation in Asparagus . . . .12 

„ in Pimis Pinaster . . . .13 

,, and spiral torsion in Asparagus (' Gard. Chron.') . 14 

„ in scape of dandelion . . . .16 

Pitcher on leaf of Pelargonium . . . .22 

Transition from flat to tubular segments of the perianth in 

Eranthis ..... 
Pitcher of Crassula arborescens (C. Morren) 
Gamopetalous corolla, Papaver bracteatum . 
Adhesion of petaloid stamen to segment of perianth, Crocus 

,, of petal, stamen and carpel, Cheiranthus Cheiri 

Synanthy, Campanula Medium (' Gard. Chi-on.') 
16. Synanthic flowers of Calanthe vestita, 
Synanthy in Digitalis purpurea (' Gard. Chron.') 

„ in Calceolaria 

20. Syncarpic apples 

Adhesion of two stems of oak (' Gard. Chron.') 
of branches of elm (' Gard. Chron.') 

,, of two roots of cai'rot, one white, the other red 

(' Gard. Chron.') 
Section through inverted and adherent mushrooms 
Bifurcated male catkin, Cedrus Libani 

„ leaf, Lamium album 








27. Bifurcated leaf. Pelargonium . . 63 

28. „ frond, Scolopendrium . . . .64 

29. Three-lobed lip of Oncidium . . .68 

30. Dialysis of corolla in Correct . . .71 

31. „ of corolla in Campanula (De Candolle) . 7 J 

32. Anomalous form of orange . . . . 7 ! 

33. 34. Disjoined carpels of orange (Maout) . . 7- r > 

35. Proliferous rose (Bell Salter) . . . .78 

36, 37. Apple flower, with detached calyx, &c. . . .79 

38. Flower of (Enanthe crocata, with detached calyx, &c. . . 80 

39. Anomalous bulbs of tulip . . . .85 

40. Displaced leaf of Gesnera (C. Moiren) . . .88 

41. Leaves of Pinus pinea . . . . .89 

42. Deranged leaves of yew . . . . 9<i 

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. ,, of natural structure in Cypripedium . . 93 

47. „ of malformed flower of Lycaste Skirmeri . . 93 

48. ,, of malformed flower of Dendrobium nobile . . 94 

49. Natural arrangement in same flower . . .94 

50. Diagram of malformed violet . . . . 'J-i- 

51. Monstrous flower of Cerastium (' Gard. Chron.') . . 97 

52. Inflorescence of Polyanthus, with tufts of leaves at the summit Ic-j 

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 .... Ill 

57. Branched inflorescence of Reseda Luteola . . . 112 

58. Thalamus of strawberry prolonged into a leafy branch ('Ameri- 

can Agriculturist ') . . . . 11 <i 

59. Flower of Verbascum, with dialysis of calyx and corolla and 

prolonged thalamus . . . . .11*; 

60. Median floral prolification of Diantlms . . .12" 

61. Leafy carpels and prolification of Danas C'i<rnf<< . 123 

62. Median floral prolification of Delpb iv iurn ,126 




Ifedian prolification, &c., «>f Orchis pyrm 

64. Proliferous rose (BeU Salter) 

65. Axillary floral prolification of Nymphaa Lotus (' < rare 

66. ,, ,, of DianQvtu 

67. Proliferous rose 

68. Proliferous rose (' Gard. Chron.') . 

69. Diagram of prolified orchid 

70. „ ,, orchis 

71. Adventitious roots from petiole of celery . 

72. Germinating plant of mango 

73. 74. Adventitious roots from leaves 

75. Hip of rose bearing leaf 

76. Leaves proceeding from the ovary of Nymph" " ep. 

77. Flower-stalk of dandelion, with leaves 

78. Tuft of leaves in place of flowers in Valeriana Bp. 

79. Scale- bearing spikelets of WiUdennvia 

80. " Rose Willow "... 

81. Viviparous flowers of Aim oivipara 

82. Formation of shoot on leaf of Episcia bicolor 

83. Adventitious buds on root of sea-kale 

84. S5. Production of adventitious bulbs in hy acini 

Chron.') .... 
86. Adventitious buds on hyacinth (' Gard. Chron ') 
87 — 92. Xepaul barley 

93. Formation of buds on fruit of Opuniia 

94. Flower-bud in the pod of Sinapis 

94*, 95. Adventitious pod in silique of Cheiranthus 
96, 97. Grapes, with adventitious fruits in interior 

98. Stamens in ovary of Bcechea diosmifolia 

99. Pollen in ovule of passion flower (S. J. Salter) 

100. Female flowers at the summit of the inflorescence 


101. Monoecious hop (' Gard. Chron.') . 

102. Superior ovary. &c., of fuchsia 

103. Hermaphrodite flower of Cariea . 

104. Ovuliferous anthers of OucurbUa sp. 

105. 106. Prolonged inflorescence of fig. 
107. Hollow turnip, with inverted leaves 



] U 




i • Gard. 


of Car ex 

18 I 





wphea mirriata (C. 


108. Normal flower of Gloxinia 

109. Regular peloria of Gloxinia 

110. 111. Structural details of erect Gloxinia 

112. Reflected corolla of azalea 

113, 114. Enlarged and erect placenta of 


115. Dimorphic leaves of Juniperus sinensis 

116. Regular peloria of Delphinium 

117. Structural details of the preceding 

118. Regidar peloria of violet 

119. Double-flowered regular violet 

120. Regular peloria of Eccremocarpus scaler 

121. . „ „ CaUleya 

122. Peloria of Calceolaria 

123. Peloric flower of Aristolochia 

124. 125. Peloric flowers of Corydalis . 

126. Rose plantain . 

127. Leafy bracts in Plantago maj jr 

128. „ scales of Dahlia . 

129. „ sepals of rose 

130. „ „ Fuchsia (' Gard. Chron.') 

131. „ calyx of primrose 

132. „ „ melon 

133. ,, sepals and petals of Geranium 

134. „ stamens of Petunia 

135. 136. Leafy anthers of Jatropha (Midler) 

137. Proliferous rose, with leafy carpels, &c. (Bell Salter) 

138. Cucumber, with adventitious leaf attached (S. J. Salter) 

139. Leafy carpels in flower of Triumfetta 

140. „ ovules of Sinapis . 

141. 142. Leafy ovules of Trifolium repens (Caspary) 

143. Portion of leafy carpel of Delphinium, with ovules (Cramer) . 

144, 145. Enlarged view of section of leafy carpel, &c, of Del- 

phinium (Cramer) . 

146. Placenta? of Dianthus, bearing ovules and carpels (' Gard. 

Chron.') ...... 

147. Ovules passing into carpels, Dianthus (' Gard. Chron.') 

148. Leafy shoot in place of ovule of Gaillardia . 


. 207 
. 207 
. 208 
. 209 




1,1st or ILLUSTRATIONS. xix 

. '!■.. PAGE 

149, 150. Leafy sepals, petals, &c, of Epllobium . . 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. Diagi'am 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. „ „ wallflower, &c. 306 

164. Passage of stamen to carpel in lily . . . 307 

165. Ti-ansition 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. ,, branch of Crataegus .... 317 

172. Spirally-twisted stem of teazel .... 321 

173. „ „ Galium . . . .323 

174. „ root (' Gard. Chron.') . . .324 

175. Interrupted growth of radish (American Agriculturist) . 327 

176. „ „ 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 brocoli (' Gard. Chron.') . . 351 

183. 184. Supernumerary leaf of elm . . . 353, 354 

185. „ „ 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. „ „ 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. „ malformed flower of Opihrys 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. „ ,, Odontoglossum Alexandra . 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-chesnut . 439 

209. Elongation of thalamus, apostasis, &c, in flower of Delphinium 

(Cramer) . 

210. Adventitious growths from cabbage leaf 

211. Crested fronds of Nephrodium molle 

212. Supernumerary petals, &.C., Datura fastuosa 

213. „ petaloid segments in flower 

214. Catacorolla of Gloxinia (E. MoiTen) 

215. Atrophied leaves of cabbage 

216. Abortion of petals, pansy 

217. Flower of Oncidium abortivum 

218. Bladder plum . 

. 441 

. 445 
. 447 
. 450 
of Gloxinia . 451 

. 452 
. 460 
. 461 
. 462 
. 464 


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 atten- 
tion of botanists, or at best as objects of mere curiosity. 
By those whose notions of structure and conforma- 
tion 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 Augustin Pyra- 
mus De Candolle that the credit is due of calling atten- 
tion to the importance of vegetable teratology. This 


great botanist, not only indirectly, but from his per- 
sonal 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 indif- 
ference, 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 con- 
tributed 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 mal- 
formation 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 

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 -auction to the doctrines 
of Goethe, and himself illustrated them by teratologics! 
observations. In France, besides important papers of 
Turpin, Geoffroy de Saint Hilaire, Brongniart, Kirsch- 
leger 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, En- 
gelmann, and Morren. A very large number of com- 
munications on teratological subjects iu the various 
European scientific publications have also been laid 
under contribution. In most cases reference has been 
given to, and due acknowledgmient made of, the sources 
whence information has been gathered. Should any 
such reference be omitted, the neglect must be attri- 
buted to inadvertence, not to design. In selecting 

1 An excellent summary of the history of Vegetable Teratology is 
given in Kirschleger's ' Essui historiqae de la Teratologic Yegetale,' 
Strasburg. 1815. 


illustrations from the immense number of recorded 
facts, the principle followed lias 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 ac- 
knowledging his obligations to the Society, under whose 
auspices this work is published, and to Mr. S. J. 
Salter, to whom the book hi some degree owes its 

The drawings, where not otherwise stated, have 
been executed either from the author's own roueli 
sketches, or from the actual specimens, by Mr. E. M. 
Williams. A larg-e number of woodcuts have also 
been kindly placed at the disposal of the author bj* the 
proprietors of the ' Gardeners' Chronicle.' 1 

As it is impossible to frame any but a purely arbi- 

1 In some instances diagrams and formulae are given in explanation 
of the conform at ion of monstrous flowers ; in general these require no 
further explanation than is given in the text, unless it he 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 

p p p p p 


shows that the sepals (s) are distinct, the petals (p) coherent, and the 
stamens (ST) adherent to the petals. 


trary definition of teratology or to trace the limits 
bel ween 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 ap- 
pearance, 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 ulti- 
mately 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 never- 
theless are very unlike in the early stages of their 
existence. 1 Following Goethe, these changes in the 
course of development are sometimes called metamor- 
phoses. In this way Agardlr admits three kinds of 
metamorphosis, which he characterises as : 1st. Suc- 
cessive metamorphoses, or those changes in the course 
of evolution which each individual organ undergoes in 

1 "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. 

5 Agardh. " Theoria Syst. Plant.," p xxiii. 


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 permu- 
tations of form and function manifested in homo- 
logous 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 organ- 
ism 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 per- 
taining 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 struc- 
ture 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 circum- 

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 carried. 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 know- 
ledge 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 organiseQl 
beings, there is a process of growth, mere increase in 
bulk as it were, and a process of evolution or metamor- 


pilosis, 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 lie 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 examin- 
ing, even though it be confessedly artificial and im- 
perfect will be better than a more philosophical 
arrangement which circumstances prevent him from 

The plan followed in this volume is a slight modi- 
fication 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. Biblio- 
graphical references and lists of the plants most fre- 
quently 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 attain- 
ment it would have necessitated for its publication a 
much larger volume than the present. 1 It is hoped 
that both the lists of books and of plants are sufficiently 
full for all general purposes. 2 

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 par- 
ticular plant named, while the prefix of the * indi- 
cates that the malformation occurs with special fre- 
quency in the particular plant to which the sign is 

Teratological alterations are rarely isolated pheno- 
mena, 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 

1 In the memoirs of Hopkirk, Kirsckleger, 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. 

2 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. 


of the several cases of malformation given in these 
pages are generally arranged according to their appa- 
rent degree of importance. Thus, in a case of prolifi- 
cation 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 prolifi- 
cation, and afterwards to those on multiplication. To 
facilitate such research, numerous cross references are 

In the investigation of teratological phenomena con- 
stant 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 nor- 
mal 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 extrava- 
gant hypothesis to surmise that this was the primitive 
structure of the flower in the higher plants. Varia- 
tions 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 re- 
producing 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. 1 

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 con- 
flicting, but it is so rather from imperfect interpretation 
than from any intrinsic worthlessness. If misused 
the fault lies with the disciple, not with Xature. 

Teratology as a guide to the solution of morpho- 
logical 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 primi- 
tive condition and gradual evolution of parts in any 

1 On this subject see a paper of M. Naudin in the ' Comptes Rendus,' 
1867, t. 64, pp. 929—933. 


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 tera- 
tological investigations and from the comparative 
study of allied forms precisely as in the case of adult 

The study of development is of the highest import- 
ance 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, be- 
cause 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 con- 
ditions, 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 classi- 
fication 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 con- 
ceptions 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 P 1 

1 It is probable that many terms and expi'essions calculated to mislead 


That teratology may Berve the purposes of syste- 
matic botany to a greater extent than might at firsi 
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 relation- 
ship existing between plants — a relationship that might 
otherwise be unsuspected. So, too, some of the altera- 
tions met with appear susceptible of no other expla- 
nations than that they are reversions to some pre- 
existing form, or, at any rate, that they are manifes- 
tations of a phase of the plant affected different from 
that which is habitual, and due, as it were, to a sort of 

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 Gasrtner, 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 Domesti- 
cation' comprise so large a collection of facts for the 

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 trn< 
condition of affairs be thoroughly well ascertained by organogenetk- 
research or other means. 


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 ob- 
served 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. 1 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 con- 
formation. Usually such changes are entirely hetero- 
morphous, and, as it were, foreign to the natural 

The practical applications of teratology deserve 
the attention of those cultivators who are concerned 
in the embellishment of our gardens and the supply 

1 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 straw- 
berry. All the fruits on the particular plants alluded to were thus 
provided as it were with a white MIL Whether this be a constant 
occurrence in the particular variety is not known. 


of our tables. The florist lays down a certain arbi- 
trary 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 in- 
elastic 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 pro- 
duce 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 'mon- 
strous ' 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 some- 
times be accounted for by that inherent tendency 
which some plants possess of exchanging an her- 
maphrodite 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, con- 




As full details relating to the disposition or arrange- 
ment 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, gene- 
rally 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 dispo- 
sition 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 : 

<->. 7 8 9 10 

] 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 


tliat the whorls alternate one with another, as may be 

represent c(l by this symbol : 

9 10 11 12 L3 14 15 16 
12 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 : 


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 arrange- 
ment 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 arrange- 


ment 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 cor- 
respond precisely with those observed in the case of 
definite and indefinite inflorescence respectively. 

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 con- 
venience 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 
Fig. 1. — Diagram stamens. Throughout this book 

showing the arrange- , . . , , 

ment of parts in a com- this arrangement is taken as the 
plete, regular, penta- stan( j ar( i f reference. Neverthe- 

merous flower: s, sepals; 

p, petals; st, stamens; less the spiral order does occur ill 

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 import- 
ance 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 develop- 
ment 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 con- 
struction 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, con- 
sidered as the primitive one transmitted by hereditary 
descent from generation to generation, and modified to 
suit the requirements of the individual, or in accord- 
ance 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 hypo- 
thetical 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 

ai;i;a\<.i:\ii;\t OF OEG \\». i 

impossible, withoul studying the development of the 
malformed (lower, to ascertain whether the altered 
arrangemenl 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 con- 
genital (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 
sime, 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, Dis- 
placement, Prolification, Heterotaxy, and Heterogamy. 



The union of parts, usually separate in their adult 
condition, is of very common occurrence as a mal- 
formation. 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 subse- 
quent coalescence of organs exist under ordinary cir- 
cumstances. Considered as a teratological pheno- 
menon, union admits of being grouped into several 
subdivisions, such as Cohesion, Adhesion, Synanthy, 
Syncarpy, Synophty, &c. Each of these sub-divisions 
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 maybe 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 (con- 
genital fusion), and that pressure will induce the 
coalition of organs primarily distinct. 



Following Augustin Pyramus 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 nnder general 
circumstances become divided during their develop- 
ment. Nothing is more common as a normal occur- 
rence, 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 

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. 
Secondhy. Two or more stems become joined together. 



Thirdly. The branches become united to the stem; 
or, lastly, the roots may become fused one with 

The first of these is most commonly met with, 
doubtless owing to the number of the branches and the 

Fig. 2. — Cohesion of two branches in Dipsacus sylvestris. 

facilities for their union. An illustration of it is afforded 
by the figure (fig. 2), showing cohesion affecting the 
branches of a teazle (Dipsacus si/lvesfris). 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, ac- 
cording to circumstances. This mode of union beloi 
perhaps, rather to the domain of pathology than of tera- 
tology. Some of the instances that have been recorded 
of very large trees. Buch as the chestnut of Mount 
.Ktna. are really cases where fusion Las taken pi 
between several of the branches, or suckers, thrown out 
from the same original stem. 1 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 rosi a, &c. 

Fasciation. — In the preceding instances of union 
between the branches, <kc, 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 

3.— Fasciation in Lettuce. 

from the softness of the vouno- tissues. In this way it 
is probable that what is termed fasciation is brought 

1 See a curious instance of this kind in the branches of JFwwa. 
'Regel. Garten Flora,' vol. 8, tab. 268. 



about. This is one of the most common of all mal- 
formations, 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. 

Fig. 4. — Fasciation in Asparagus. 

Very generally the surface is striated by the pro- 
minence 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.^.in the 
ash ( Fraxinus), wherein it has been likened to a shep- 
herd'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 

PlG. 5. — Fasciated branch of Plans Pinaster 



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 sub- 
divisions. These latter maybe deformed like the parent 
branch, or they may resume the ordinary aspect of the 

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 



Fig. 6. — Fasciation and spiral torsion in the stem of Asparagus 


orest along 1 the apex. When, as often happens, the 
deformity is accompanied witli 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 
Bpecimen of Bwplev/rum falcai/wm 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 flat- 
tened, and the different organs of the flower are arrange* I 
on an elliptical axis. A case of this nature is described 
by Schlechtendal (' Bot. Zeit.,' 1857, p. 880), in Oytisus 
rrigricans, 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 hyper- 
trophied and fasciated, and bore on its surface a few 
adventitious buds. 1 The pedicels of Sbreptocarpus Bexii 
have also been observed in a fasciated state. 2 

It has been occasionally observed that the fasciated 
condition is hereditary ; thus, Moquin relates that 
some seeds of a fasciated Oi/rsiv/m reproduced the same 
condition in the seedlings, 3 while a similar tendency 
is inherited in the case of the cockscomb (Gelosia). 

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 men- 
tioned, Moquin was of opinion that fasciation was due 
to a flattening of a single stem or branch. Linnams, 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 dicisoletjplanta eumplures ccmles 
connascvntiif, ut unus ex phirimis instar fascice eva<lif 

1 'Bull. Soc. Bot. France,' I860, p. 881. 
■ Ibid., 1861, p. 708. 
3 Ibid.. 1860, p. 923. 



et compressus" (Linn., ' Phil. Bot.,' 274). A similar 
opinion was held by J. D. Major in a singular book 
entitled ' De Planta Monstrosa, Gottorpiensi,' Schles- 
wig, 1665, wherein the stem of a Chrysanthemum is 
depicted in the fasciated condition. 

The striaa, which these stems almost invariably present, 
exhibit the lines of junction, and the spiral or other cur- 
vatures 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 tog-ether. Prof. Hincks 1 cites cases of this kind 

Fig. 7. — Fasciation in the scape of the Dandelion (Leontodon Taraxacum). 
1 ' Proc. Linn. Soc.,' April 5, 1853. 


in Primula vulgaris, Hierackim aureum,a,nd Rantmeulua 
bulbosus. 1 have myself me1 with several cases of the 

kind in Primula verts, in the Polyanthus, in the Daisy, 
and in the Leontodoti 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 par- 
ticular 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 oo, 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 re- 
sembling the figure of oo, but when more completely 
combined they have an outline of a very elongated 
figure, and something similar is to be expected in her- 
baceous stems. Even the elongated pith of a trans- 
versely 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 

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 con- 
sidered 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 trans- 
verse 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 1 
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 nourish- 
ment, 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. Secki/m cristatwm, 

1 'Organ. Voget.,' pi. iii, fig. 1. 


Gelosia, &c. This condition may be induced by the 
art of the gardener — " Fit idem arte, si pl/ures caules 
enascentes cogani/wr penetrare coarctatwn spati/wm et par- 
turiri tanquarn ex.cmgvsto utero, sic scepe im Uammculo, 
Beta, Asparago, Hesperide I'i'iiv, ('closid, Tnnjopogone, 
Scorzonsrd (Jotitla faiidit" Linnasus op. cit. 

Plot, in his ' History of Oxfordshire,' considers fas- 
ciation to arise from the ascent of too much nourish- 
ment 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 
©f 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 applica- 
tion 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. 




a. Herbaceous. 

Ranunculus tripartitus. 
bulbosus ! 
Delpbinium elatum. 
* sp. ! 

Hesperis naatronalis. 
*Cheiranthus Cheiri ! 
*Matthiola incana ! 
*Brassica oleracea ! var. pi. inflor. 
Linum usitatissimuni ! 
Althaea rosea ! 
Lavatera trimestris. 
Geranii sp. 
Tropseolum majus ! 
Viola odorata inflor. ! 
Reseda cdorata ! 
Fragaria vesca. 
Ervum lens. 
Trifolium resupinatum. 

repens ! 

pratense ! 
Saxifraga mutata. 

Bupleurum falcatum. 
Bunium flexuosum. 
*Sedum reflexum ! 
cristatum ! 
Epilobiuni augustifolium ! 
Momordica Elaterium ! 
Gaura biennis. 
Cotula foetida. 
Barkhausia taraxacifoHa. 
Carlina vulgaris ! 
Apargia autuninaHs. 
*Leontodon Taraxacum inflor. ! 
Centaurea Scabiosa. 
*Cichoriuin Intybus! 
Hieraciuni Pilosella. 
*Chrysanthemurii Leucantbemum. 

indicum ! 
Antbemis nobilis. 

Cirsiuru lanceolatuni. 
Conyza squarrosa ! 
Inula dysenterica ! 
Tragopogon porrifolium. 
Cnicus palustris. 
Carduus arvensis ! 
Heliantbus tuberosus ! 

Cineraria palustris. 
HeHanthus sp. ! 
Dahha variabilis. 
Bellis perennis inflor. ! 
Coreopsis sp. ! 
Crepis virens. 
Lactuca sativa ! 
Zinnia elegans. 
* Campanula mediuni ! 

Dipsacus pilosus. 


Knautia arvensis. 
Pbyteuma orbiculai'e. 
Jasione montana. 
*Linaiia pui*purea ! 
Antirrhinum majus ! 
Veronica amethystea. 
Veronica niaritima. 

RusselKa juncea ! 
Digitalis purpurea ! 
Ajuga pyramidalis. 
Hyssopus oflBcinalis. 
Dracocephalum moldavicum. 
Myosotis scorpioides. 
Echium pyrenaicum. 

Stapelise sp. 
Lysimachia vulgaris ! 
Androsace maxima. 
Primula veris inflor. ! 

denticulata inflor. ! 
Polemonium cceruleuni. 
Convolvulus sepium ! 
ai*vensis ! 
Plantago media. 
^Euphorbia Characias. 
Suceda niaritima. 
*Celosia sp. 
Beta vulgaris inflor. ! 
Phytolacca sp. 

(3. Woody. 

Berberis vulgaris. 
Hibiscus syriacus ! 
Acer pseudo-platanus ! 
Dodoncea viscosa. 
Stercuba platanif oba. 
Euonymus japonicus ! 
Vitis vinifera inflor. ! 



Spartium Seopariuni ! 
Spartium junceuru ! 
Cytisus Laburnum. 

Cborozema ilicifolinm. 
Amorpha sp. 
Phaseolus sp. 
Primus sylvestris. 

Laurocerasus ! 
Rosa sp. ! 
Spiraea sp. ! 

Cotoncaster microphylla ! 
Ailanthus glandulosus. 
*Fraxinus Ornus ! 

excelsior ! 
Melia Azedarach. 
Xanthoxyluni sp. ! 
Sambucus nigra ! 
Aucuba japonica. 
Erica sp. cult. 
Jasminum nudiflorum ! 

officinale ! 
Olea europcea. 
Punica Granatum. 
Ilex aquifolium ! 

Dapbnc indica. 
Dapbne odora. 
Suasda fruticosa. 
Ulmus campestris. 
Alnus incana. 
Sabx vitellina, &c. ! 
Thuja orientalis. 
Pinus pinaster ! 

sylvestris ! 
Abies excelsa ! 
Taxus baccata. 
Larix europcea. 


Lilium Martagon. 
candidum ! 
*Fritillaria imperialis ! 
Asparagus officinalis ! 
Hyacintbus orientalis ! 
Tamus communis ! 
Narcissi sp. ! 
Gladiolus sp. 
Zea Mays. 

See also — Moquin-Tandon, ' Elem. Ter. Veget.,' p. 146 ; C. O. Weber, 
1 Verhandl. Nat. Hist.,' Vereins, f. d. Preuss., Rbeinl. und Westpbal , 
1860, p. 347, tab. vii; Hallier, ' Pbytopatbol.,' p. 128; Boebmer, '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. Tilice,' Berlin, 1861.) It is also stated 
that this condition is not perpetuated by grafting. 

I have in my possession a leaf of Antirrhinum majus, 
and also a specimen of Pelargonmm, wherein the blade 

Fig. 8. — Pitcher-shaped leaf of Pelargonium. 

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 
PelargonwTn p&ltatwm with those of P. cucuMabwm ('Cav. 


Diss.,' tab., L06) will sliow how easy I lit- 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 1 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 a notices something of the same kind in 
the leaf of Amorpka fruticosa; Treviranus 3 in that of 
. hrislolochia Si pho. 

M. Puel 4 describes a leaf of Poly gonatu/m multifiorum, 
the margins of which were so completely united to- 
gether, 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 As- 
sociation 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, Trifolwm r&pens, 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 Mcw- 
ijriiiirin, which appear to originate from the union of 
the margins of the reflected leaf. 

Tubular petals occur normally in some flowers, as 
Helleborus, Epwrndw/ni, Viola, &c, and as an excep- 
tional occurrence I have seen them in Ranunculus r&pens, 
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 Sandy well 
Park, I am indebted for the flower of a Pelargonium, 

1 ' Bull. Acad. Roy. Bruxelles,' t. xviii, p. i and p. 591. 

2 ' Linnaea,' torn. 13, p. 383. 

3 ' Verhandl. Nat. Hist. Vereins,' 1859, Bonn, torn, xvi, tab. 3. 

4 ' Bull. Soc. Bot. Pr.,' vol. i, p. 62. 



in which one of the petals had the form of a cup sup- 
ported on a long stalk. This cup -shaped organ was 
placed at the back of the flower, and had the dark 

Fig. 9. — Eranthis hyemalis. Transition from flat sepal to tubular petal. 

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 re- 
semble perfect corollas. 

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. 1 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* but as a similar con- 
dition 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 

1 ' Report of Internat. Bot. Congress,' London, 1866, p. 131, tab. 
vii, figs. 10 — 13. 

2 ' Bull. Acad. Roy. Belg.,' t. xviii, 2nd part, p. 179. 


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 ex- 
pansion, 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 pre- 
ceding, 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 Tropceotum 
ma/jus, and Strelitzia regma; 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, oxyph/ylla? M. Clos 3 has ob- 
served 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. 3 

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 pertmum, as well as some cases of a 
similar kind that are occasionally met with, as instances 

1 D. C, ' Organ. Veget.,' pi. xvii, fig. 3, and pi. xlviii, fig. 2. 

2 ' Mem. Acad. Toulouse,' 1862. 

3 Bonnet, ' Recherches Us. feuill.,' pi. pa, fig. 2. 



of the cohesion of the margins at the base and apex 
of the leaf, which thus appears perforated. This ap- 
pearance, 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 occasion- 
ally takes place, as in Eberms cretiGa. 1 It has also 
been observed in Tithonia, and is of constant occur- 
rence in the seed leaves of some MesembryantTiema. 
This condition must be carefully distinguished from 
the very similar appearance produced by quite a dif- 
ferent 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 Crasmla a/rborescens, observed by C. 

Fig. 10. — Two-leaved pitcher of Crasmla arborescetis, after C. Morren. 

The stipules may also be fused together in different 

1 De Candolle, ' Mem. Leg.,' pi. v, fig. 14. 


wnvs: their edges sometimes cohere between the leaf 
and tlif stem, and thus form a solitary intra-axillary 
stipule. At other times they become united in such a 
manner as to produce a .-ingle 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 speci- 
mens 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 Qypripedvum, &c. An illus- 
tration of this occurrence is given by Mr. J. T. 
Moggridge in Oph/rys msecttf&ra 9 m ' 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 la- 
bellum, dislocation of the parts of the flower, &c. 

Cohesion of the petals. — Linnasus mentions the occur- 
rence of cohesion of the petals in Scvponaria. 1 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 

Fig. 11. — Ganiopetalous flower of Papaver bradeatum. 

irregular one, like the corolla of Labiates, is recorded 
by Jaeger. 2 There is in cultivation a variety of 
Pa-paver bracteatum, in which the petals are united by 
their margins so as to form a large cup. Under 

1 ' Phil. Bot.,' § 125. 

2 ' Nov. Act. Acad. Nat. Cur.,' 14, p. 642, t. xxxvii. 


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. In- 
dications 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, Poly- 
adelphia) ; at other times, by the cohesion of the 
anthers (Syngenesia), in which latter case the union is 
generally very slight. It must be remembered, how- 
ever, 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 maybe 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. 1 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 oc- 
currence in plants, under ordinary circumstances, but 
is less commonly met with than might have been ex- 
pected as a teratological phenomenon. 

1 ' Journal of the Linn. Soc. Bot.,' vol. iv, p. 55. 


Further details relating to cohesion of the various pai*ts of the flower 
are cited in Moquin-Tandon, ' El. Ter. Veg.,' p. 248 ; ' Weber. Verhandl. 
Nat. Hist. Yereins f. d. Preuss. Rheinl. und Westphal.,' 1860, p. 332, 
tabs. 6 et 7. 

Formation of ascidia or pitchers. — In the preceding para- 
graphs, 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 separa- 
tion 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 Pro- 
fessor Charles Morren, 1 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 com- 
prises 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 N&penthes, &c, which occur normally 
and constantly, are not here included. Possibly some 
of the cases would be more properly classed under 
dilatation or excavation. 

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 europsea ! 

Pelargonium inquinans ! 

Staphylea pinnata. 

Amorpha fruticosa. 

Pisum sativum ! 

Lathyrus tuberosus. 

Vicia sp. 

Gleditschia sp. 

Ceratonia siliqua. 

TrifoHum repens ! 

Cassia marylandica. 

1 • BuU. Acad. Roy. BrnxeUes,' 1838. t. v. p. 582. ' Bull. Acad !:• 3 
Belg.,' 1852, t. xix, part hi, p. 437. 


Mimosa Lophantha. 
Rosa oentifolia. 

Begonia sp. 
Bellis perennis ! 
N tiana sp. 
Goodenia ovata '. 
Antirrhinum majus ! 
Tinea roe 

Polygonum oriental.;. 
Aristolockia sipho F 

:;rum varieu'atum var. .' 
Spinaeia oleracea. 
Corylus avellaua ! 
Polygonatum miiltiflornm. 
Xanthosoma appendiculatum ! 

2. Calyptriforni or hood-like pitcher?, formed by the complete union 
of the margins, and falling off by a transverse fissure (as in the calyx ol 

Tuhpa Gesneriana. 

B. Pobjphyll 

1. Diphyllous. formed by the union of two leaves into a single cap, 

tube, or funnel. &e. 

Pisum sativum, stipul - 
Crassula arborescens. 
Polygonatum multifiorum. 

2. Triphyllous, formed by the union of three lea - 
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 fullowins treatises on the subject of ascidia : — Bonnet, 
• Rech. Us. Feuilles,' p. 216, tab. xxvi. f. 1, Brassica ; De Candolle, 
'Trans. Hort. Soc.,' t. v, pi. 1. Brassica; Id., ' Org. Veget,' I. 316; 
'Bull. Soc. Bot. Ft.,' I, p. 62, Polygonatum ; -Bull. Aead7Belg.. ; 1851, 
p. 591, Rosa ; Hoffmann, ' Tijdschrift v. Xatuur. Gesehied.." vol viii, 
p. 318, tab. 9. Geratonia; C.Mulder. • Tijdschrift. Sua., vol. vi. p. 106, 
tab. 5, 6, Tr [folium , Mimosa, Staphylea ; ' Mblkenboer,' p. 115. t. 4. 
Li sica. 




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 con- 
sequence 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 neces- 
sarily 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 ordi- 
nary circumstances in the bract of the Lime tree, which 
adheres to the peduncle, also in Neuropdtis, while in 
Erythrockitcm liypophijllanthus the cymose peduncles are 
adherent to the under surface of the leaf. 

Adhesion between the axes of the same plant is suffi- 
ciently treated of under the head of Cohesion, from 
which it is in this instance impossible to make a dis- 
tinction. 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 

01 i.i-:\ 

it generally forms but a pari of other and mi 
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 Agavt 
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, 

mera, fyc} 

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 adh-esion 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 ter- 
minal 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. 8 The same author mentions having seen a 
remarkable example of adhesion in the involucels of 
Gaucalis leptophylla, the bracts of which were soldered 
to the outer surface of the flowers. M. Bureau 3 men- 

! "Wydler, ' Flora,' 1852, p. 737, tab. ix. 

- • El. Ter. Veg.,' p. 254. 

3 ■ Bull. Soc. Bat. Fr..' 1857, p. 451. 



tions an instance wherein the spathe of Narcissus bi- 
fiorus 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 sur- 
faces 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 con- 
siderable 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 byMorren 
as calyphyomy, koXvZ tyvo/iiai. 1 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 Catthya 
Forbesii was adherent to the labellum. 2 

Adhesion of the stamens to the petals is of common occur- 
rence under natural circumstances. Cassini has de- 
scribed a malformation of Centaurea coll inn, 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 infre- 

1 ' Bull. Acad. Belg.,' vol. xix, pai-t ii, p. 335. 
? ' Bull. Soc. Bot. Fr..' 1860. p. 25. 



quenl occurrence among Orchids. I have observed 
cases of the adhesion of the segments of the perianth to 
the stamen in Oph/rys a/ranifera, Odontogh8Svm t sp. 

&c. It is the ordinary condition in 
Gongora and some other genera. I 
have seen it also in Lilmm lancifoli/wm. 
Some forms of Omen*, occasionally met 
with, present a very singular appear- 
ance, 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 1 speaks of a similar union in 
Tigridia pavonia. Morren 2 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. FlG 12 _ crocu*. 
It should be remarked that it was Adhesion of petaloid 
the outer series of stamens that were stamens to perianth. 
thus fused. 3 

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 cmstrale* 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, compara- 
tively 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 

1 ' Bull. Soc. Bot. Fr.,' 1861, p. 147. 

2 * Bull. Acad. Belg.,' vol. xviii, part ii, p. 498. 

3 See also Pvillieux, ' Bull. Soc. Bot. Fr.,' 1861. p. 195. 

4 ' Mem. Acad. Toulouse,' 5th series, vol. iii. 



lip of the corolla of Salvia pratensis. 1 In the ac- 
companying 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 impres- 
sion 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. 

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 swperus" fre- 
quently becomes inferior from de- 
tachment from, or from want of 
union with the surface of the ovary. 

Adhesion of fruit to branch. — Of this Mr. Berkeley 2 cites 
an instance in a vegetable marrow {Cum/mis), 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 

1 Linnsea,' vol. ii, p. 607. 

2 ' Journal Roy. Hort. Soc.,' new ser., vol. i, 1866, p. 200. 

Fig. 13. — Cheiranthns 
cheiri. Adhesion of 
petal to stamen and 
open carpel. 

M \ WTIH . 


arc united together, so that we have a single peduncle, 
bearing at its extremity two flowers placed in approxi- 
mation 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 Umbelliferce. Union of 
this kind occurs frequently in the common cornel 
{Coram), 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 Cent a urea. In other cases the 
union involves not only the stalk but the flowers them- 
selves ; thus fusion of the flowers is a common accom- 
paniment of fasciation, as was the case in the Campa- 
nula figured in the cut (fig. 14). 

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 incorpo- 
rated 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, tiymphyo- 
myrtus, &c, while the large size of some of the culti- 
vated 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, 1 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 ckrysanthus. 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 Aristolocliia Clematitis? 

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 Nar- 
cissus incomparahilis I remarked a ten-parted perianth, 

i ' Bull. Soc. Bot. Fr..' 1861, p. 159. 
: Ibid.. L859, ]». 167. 

S3 vwiiiy. 


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. 

As an illustration of a more complicated nature 
reference may be made to three flowers of Aconitum 
Na/pellus, figured by A. de Chamisso, ' Linnsea,' 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 

Fig. 15. — Union of three flowers Fig. 16. — Shows the abortion of 

of Calanthe vestita. the central spur in synanthic 

flowers of Calanthe vestita. 

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 flow^ers is occupied by a two to 
five-celled pistil, between the carpels of which, not un- 
frequently, the stem of the plant projects, bearing on 
its sides bracts and rudimentary flowers. (See Proli- 

Fig. 17. — Syrianthy and other changes in a Foxglove. 

fication.) An instance of this nature is figured in the 
' Gardeners' Chronicle,' 1850, p. 435, from which the 
cut (fig. 17) is borrowed. 

One of the most singular recorded instances of 
changes connected with fusion of the flowers is that 



cited by Reinscb, 1 where two female flowers of SaUx 

ciiicmt 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 ob- 
served, it is only necessary to 
allude to a few of the most salient 

Sometimes the upper lips of two 
flowers are fused into one, the two 
lower remaining distinct. In other 
cases, the upper lip disappears 
altogether, Avhile there are two 
lower lips placed opposite one 
another; of the stamens, some- 

Fig. 18. — Synanthic 
flowers of Calceolaria in 
which, with tAvo upper 
lips, there was but a 
single lower one. 

1 ' Flora,' 1858, p. 65, tab. ii. 


times the outermost, at other times the innermost 
disappear. 1 

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 anc&ps, 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. Mog- 
gridge 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 
Gentranthus 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 2 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 

1 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. 

2 ' Bull. Soc. Bot, Fr.,' vol. vii. p. 625. 


as to form but a single tube. It is not uncommon, 
as lias been before stated, to find two corollas enclosed 
within one calyx, but this is probably 1 lie 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 l than where 
they are remote ; so too, the upper or younger por- 
tions of the inflorescence are those most subject to 
this change. In like manner the derangements con- 
sequent 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 resist- 
ing the effects of that agency, than they are in the 
outer portions of the flowers where growth is less 

Morren in his papers on synanthic Calceolarias, 
before referred to, considers that the direction in 
which fusion acts is centripetal, e. g. from the cir- 
cumference 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 

1 Cramer, ' Bildungsabweichungen,' p. 56, tab. vii, fig. 10, figures a 
ease wherein the two central flowers of the eapitulum of Centaur ea Jacea 
were united tog-ether. 



is reversed. This remark applies particularly to indi- 
vidual 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 par- 
ticular 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 aurantiuui. 
*Fuchsia var. hort. ! 
CEnotbera sp. 
Saxifraga sp. 
Podalyria niyrtillifolia. 
Prunus Arrueniaca. 

Pyrus Mams. 
Persica vulgaris. 
Crataegus monogyna. 
Robinia pseudacacia.' 
Gleditschia triacantbos. 
Syringa persica. 
Cornus sanguinea. 
Viburnum sp. 
*Lonicera sp. plur ! 
Centrantbus ruber ! 
Valaiitia cruciata. 
Centaurea moscbata. 

Zinnia elegans. 

Zinnia revoluta. 
Heliantbus sp. ! 
Spilantbes oleracea. 

# Leontodon Taraxacum ! 
Senecio Doria. 
Cicborium Intybus. 
Lactuca sativa. 
Antbemis retusa. 
*Campanula medium ! 

Azalea indica ! 
Vinca minor. 
Atropa Belladonna. 
*Solanum Lycopersicuni 

* Petunia violacea ! 
Galeopsis ochroleuca. 
Betonica alopecuros. 
*Digitalis purpurea ! 

* Antirrhinum majus ! 
*Linaria purpurea ! 

!lc Pedicular is sylvatica ! 
*Calceolaria var. hort. ! 
Scrophularia nodosa. 
Salpiglossis straminea. 
Streptocarpus Rexii. 
*Gesnera var. hort. ! 
JEschynanthus sp. ! 
Tbyrsacanthus rutilau • ! 
Anaprallis colliua. 

M \r\i;n . 


■ Primula veris ! 

*Primnla acaulis, var. umbellata! 

elatior P 

* sinensis! 

A rist i >1< M-hia Cleinatitis. 
Blitum sp. 
Chenopodium sp. 

KlllIH'\ S]>. 

Sails cinerea. 

*Hyacinthus oriental is ! 
Lilium bulbif emm ! 

croceum, et sp. alix, pi. 
Tnlipa, sp. 
Polygonatum anceps. 

Fritillaria imperialis ! 

A^iw americana. 
his versicolor. 

Crocus, sp. 

Colehicum autumnalc. 
Narcissus incomparabilis ! 



*Opbrys aranifera ! 
Calantke vcstita ! 
Oncidium bicolor. 

&c. &c. 

In addition to tbe works before cited, additional information on tbis 
subject may be gained from tbe following : — Jaegei*, ' Missbilld.,' p. 92. 
v. Scblecbtend, ' Bot. Zeit.,' 1856, Robinia. Weber, ' Verbandl. Nat. 
Hist. Vereins. Preuss. Rbeinl.,' 1849, p. 290, Primula. Hincks, ' Rep. 
Brit. Assoc. Newcastle.' l*:'>s, Salpiylossis. Clos, ' Mem. Acad. Toulouse,' 
vol. vi, 1862, Anagallis. Wigand, 'Flora,' 1856, tab. 8, Pedicularis. 
Henfrey, ' Botan. Gazette,' i, p. 280, Reseda. P. Reinscb, ' Flora,' 1860, 
tab. 7, Petasites. Weber, ' Verbandl. Nat. Hist. Vereins. f . d. Preuss. 
Rbeinl. u. Westpbal.,' 1860, p. 332, tabs. 6 et 7, Primus, Persica, Cam/pa- 
nula, Taraxacum, Saxifraga, Silene, Hyacintlms, &c. Miquel, ' Linnaea,' 
xi, p. 423, Colehicum. Micbel, ' 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 altera- 
tion in the other parts of the flower, as happens 
normally in many Ga/prifoliacece, Bubiacece, &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 inflo- 
rescence 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 con- 
dition, 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 indi- 
vidual 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, goose- 
berries, 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 Rawimcidus tri/partitus 
soldered together. He has also seen three melons 
similarly joined. 1 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. 2 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. 3 

1 ' Bull. Bot.,' tab. iii. figs. 4—6. 

s 'Mem. greffe Ann. Science Nat.,' ser. i, t. xxiv, p. 334. 
3 "Mespilus portentosa." Poit. et Turn., ' Poinol. Franc.,' liv. xxxi. i>. 
202, pi. 20 -1. 

81 Nl A.RPY. 


A very remarkable example of Syncarpy lias been re- 
corded by E. Koeniof is which nine strawberries were 

■ CD 

borne on one stem ( Wragaria bot/ryformis), 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 

•■ 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. 

In many of these cases, where the fruits are united 

Fig. 19. — Adhesion of two apples. 

Fig. 20.— Section of united apples. 

by their bases, the summits become separated one from 
the other, so as to resemble the letter V. Such di- 

1 Duchesne, ' Hist. Nat. Frais..' p. 79. 


vergence 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 inter- 
penetration 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. Rceper has observed two apples grafted together, 
one of which had its stalk broken, and seemed evi- 
dently borne and nourished by the other apple ; l 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 pre- 
cisely similar appearances. 2 

Fusion of two or more nuts (Gorylus) is not un- 
common ; I have seen as many as five so united. 3 In 
these cases the fruits may be united together in a ring 
or in linear series. 

In some Leguminosce, 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 Gcesabpinia 
digyna this is so commonly the case as to be consi- 
dered almost the normal state. (De Cand. ' Mem. 

» De Cand., ' Pliys. Veget,,' torn, ii, p. 7S1. 

2 Sched. de monstr. plant, ' Act. Helv.,' tab. i, fig. 8. 

3 ' Mem. gveffe,' loc. cit., tab. xxiv, p. 334. 

SYNI MM'V. 1'.' 

I. j-.,' pi. 2, fig. G; pi. 3, fig. 2.) At times tl 
carpels become fiised together, and it becomes difficult, 
whni 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 synearpy is 
difficult to distinguish. It may, nevertheless, be said as 
a general rule that the union brought about bv 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 

An analogous phenomenon takes place not uncom- 
monly in mosses, the spore capsules of which become 
united together in various ways and degrees. Schimper 1 
cites the following species as subject to this anomaly : — 
Buxbaumia indusiata, Leskea sericea, Hypnum lutesd s, 
Anomodon altematus, Glinadum dendroides, Bryum ccespi- 
titium, Brachythecium plumosum, Mnvum serratum, 
Splachnum vasculosum. It has also been observed in 
Trichostomum rigiekdum and Hypnum triquetrum. 

In addition to the authorities already mentioned, the reader may 
consult Moquin-Tandon. ' El. Ter. Teg.,' p. 270. Turpin, ' Mem. greffe. 
Ann. Sc. Nat.,' ser. i, t. xxiv. p. 334. De Candolle. ' Organ. Vegefc.,' t. i. 
Duhamel, ; Phys. des Arbres,' t. i, p. 304, tab. xiii, xiv. Weber. 
'Verhandl. Xat. Hist. Yereins f. d. Preuss. Rheinl. u. WeatphaL,' 
I860, p. 332, tab. vi et vii. 

' ' Bull. Soc. Bot. Franc.,' 8, pp. 73 and 351. tab. ii ; and Rose. ' Bot.' x. p. 410. 



Synspermy, or Union of the Seeds. — Seeds may be united 
together in various degrees, either by their integuments, 1 
or by their inner parts. Such union of the seeds, 
however, is of rare occurrence. It takes place nor- 
mally, 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 pro- 
portion as the contact between the younger growing 
portions of the two axes respectively is close. The 
huo-e size of some trees has been, in some cases, attri- ■ 
buted to the adnation of different stems. This is said 
to be the case with the famous plane trees of Bujuk- 
dere, near Constantinople, and in which nine trunks 
are more or less united together. 3 

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 

1 Nijmplicm lutea, JExcnlus Hippocastanum, &c. See Moquin, ' El. Tor. 
Veg.,' p. 277. a 0. Marlins. 'Promenade Botanique,' p. 8. 



(fig. 2-*, A), with its small overgrown pari above the 

junction, took the colour and large dimensions of the 
white Belgian (</), which, in like manner, with its 

Fig. 21. — Adhesion of two distinct stems of oak. or possibly cohesion 
of branches of the same tree. ' Gard. Chron.,' 1846, p. 252. 

larger head above the joining (a), took the colour and 
small dimensions of the red one at and below the 
union (e '7). The respective qualities of the two roots 



were thus transposed, while the upper portions or 
crowns were unaffected : the root of one, naturally 

Fig. 22. — Adhesion of the branches of two elms. 
1849, p. 421. 

Gard. Chron.,' 

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) . 

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 stumpjRepends on similar 
facts. In Abies pectmata, says Goeppert, 1 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. 

1 ; Ann. So. Nat./ t. xix. 1843, p. 111. tab. iv. 



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 in- 
verted position on the top of its fellow. 1 

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 

1 'Ann. Nat. Hist.,' ser. 2, vol. ix, tab. xvi. 
p. 352, &c. 

Phytologist,' 1857, 


induced artificially ; thus, the employment of the root 
as a stock, " root-grafting," is now largely practised 

Fig. 24. — Section through two adherent mushrooms, the upper one 

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. 1 

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 

» Quoted from the • Revue Hortic' in ' Gard. Chron.,' 1866, p. 386. 


blue, while Borne produced tubers partly white and 
partlv blue. 

Adhesion of the axes of plants belonging to different species 
is a more Bingular 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 essen- 
tial 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 bo- 
tanical 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 (Viscu n), which may be found growing on 
plants of very varied botanical relationship, the occur- 
rence of occasional adhesion between plants of distant 
affinity is not so much to be wondered at. Union be- 
tween the haulms of wheat and rye, and other grasses, 
has been recorded. 1 Moquin-Tandon ~ relates a case 
wherein, by accident, a branch of a species of Sopl 
passed through the fork, made by two diverging 
branches of an elder (Samlmcus), growing in the 
Jai'din des Plantes of Toulouse. The branch of the 
hora contracted a firm adhesion to the elder, and 
what is remarkable is that, although the latter has much 

1 Senebier, ' Phys. Veget., 1 t. iv, p. 426. The same author also cites 
Romer as having found two plants of Ramuneuhts, from the stem of 
•which emerged a daisy. As it is not an uncommon practice to stick a daisy 
• >n a buttercup, it is to be hoped no hoax was played off on M. Romer. 

- ' El. Ter. Ye?./ p. 289. 


softer wood than the former, yet the branch of the 
harder wooded tree was flattened, as if subjected to 
great pressure. 1 It is possible that some of the 
cases similar to those spoken of by Columella, Virgil, 2 
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 
Cycadece, Coniferce, &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 

1 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 ! 

2 See Daubeny, ' Lectures on Roman Husbandry,' p. 156. 

svNoPim. 57 

of their axes, leaving merely the four cotyledons sepa- 
rate. A similar actuation has been observed by the 
same botanist in Lepidvu/m satwv/m and Swia/pis ramosa, 
as well as in other plants. 1 I have met with correspond- 
ing instances in Antirrh/vmwn ma/jus 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 " rhizo- 
collesy" to this union of the roots. 2 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 ferti- 
lisation of Fuchsia cocci uea (quere F. magellanica ?) by 
the pollen of F. fulgens. 

1 A. P. De Candolle, ' Organ Veget.,' t. ii. p. 72, tab. liv, fig. 1. 
- ' Bull. Acad. Belg.,' t. xx, part i, 1852, p. 43. 



Uxdee 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 in- 
cludes 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 in- 
tegrity 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 load 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 
w^ell of the ordinary as of the unusual course, of deve- 
lopment in any particular flower is thus required in 
order to ascertain with accuracy the true nature of the 
separation of parts. The late Professor Alorren 1 pro- 
posed the general term Monosy (/uo'vwtnc) for all these 
cases of abnormal isolation, subdividing the group 
into two, as follows — 1, Adesmy (a-Sfoyxoe) , including 
those cases where the separation is congenital ; and 
2, Dialysis (SmAu'w), comprising those instances where 
the isolation is truly a result of the separation of 
parts previously joined together. Adesmy, moreover, 

> ' Bull. Acad. Belg.,' t. xix.. part iii, 1852, p. 315. 


was by the Belgian .-avant said to be homologous 
when it occurred betwe< u members of the same whorl, 
e.g. between the sepals of an ordinary monosepali 
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 Dial- 
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. 



Whhn 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 appro- 
priate one to be applied to the subdivision of an habi- 
tually 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. 1 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 2 
gives as an illustration the case of a specimen of 
Rusci's 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 marsin, the notch beingr 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 Intra, 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 

i Loc. cit.. p. 295. 

2 'Bull. Soc. Bot. France,' 1857, p. 758. 

I [SS10N. 


;ilso seen it iii Plantago lanceolata, Reseda hiteola, 
Campanula medium, Epacris i/mpressa, and a bifurcation 
of the axis of the spikelel within the outer glumes in 
Lolium perenne* and Anthowiithiihi <><h>f<ititni. In bhe 
Kiw Museum is preserved a cone of Abies exceha* 
dividing into two divisions, each bearing bracts and 
scales. A similar thing frequently occurs in i he male 
catkins of Ced/rus Libani (fig. 25). 

Fig. 25. — Bifurcated male inflorescence, Cedm.s 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. 

Fission of foliar organs. — Many leaves exhibit constantly 
the process of fission, such as the SaKsburia adiantifolia, 
and which is due perhaps as much to the absence or rela- 
tively 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 

1 Masters, ' Jourl. Linn. Soc' vol. vii. p. 121. 

' Cramer, ' Bildungsabweichungen,' p. 4. tab. vi. fig. 4. figures a case 
of ibp same kind in Pinus Cerribra, 



Merciirlalis in which the leaves were deeply slashed. In 
Chenopodmm Qwinoa the leaves were so numerous and 
the clefts so deep, that the species was hardly recognis- 
able, while on a branch of Rims t 'otirms observed by De 
Candolle the lobes were so narrow and so fine as to 
give the plant the aspect of an Umbellif&r. Wigand 
('Flora,' 1856, p. 706) speaks of the leaves of Dipsams 
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 Scdbiosa 
abropwrpurea 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. 1 Steinheil has also recorded a Cern*- 
tium 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, Ukrvus 

Fig. 26. — Bifurcated leaf of Lamium album, &c. 

camppztriz, audalso in the common nettle, Urtica dimca, 

1 'Ann. des Science Nat .' 2nd scries, t. iv, p. 147, tab. v, figs. 3 and 4. 

i [8SI0S 01 leaves. 63 

the Leaves of which latter thus resembled those of Urtica 
biloba, which are habitually bilobed at the summit. M. 
Clos 1 mentions an instance where the terminal leaf and 
first bract of Orchis sa/mbucina were divided 'into two 

segments. The same author also mentions the lea 
of An&mdopsis 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 fit' cleavage. Fig. 26 represents a case of this 
kind in Lamiwm album, conjoined with suppression of the 
flowers on one side of the stem. I have also in my her- 
barium a leaf of Arwm maculatwn^ with a stalk single at 
the base, but dividing into two separate stalks, each bear- 
ing 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 Pelar- 
gonium presented the same appearance. 

Fig. 27.— Bifurcated leaf of Pdargondwm. 

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 

' 'Mem. Acad. Scien. Toulouse,' oth seine-. voL Hi. 



of France (April 3rd, 1858) exhibited a leaf of Gerasus 
Lauro-Cerasus divided in such a manner as to resemble 
a leaf of Citrus or of PJvylla/rthron. In this case, there- 
fore, the disunion must have taken place laterally, and 

Fig. 28. — Bifurcated frond, Scolopendrium vulgare. 

not from apex towards base, as is most common. 
The leaves of the common horseradish, GocMearia 
Armoracia, are very subject to this pinnated subdivision 
of the margin, and numerous other illustrations might 
be given. 

A. Braim 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 re- 
maining entire. (Verhandl., d. 3-5, Naturvorschef- 
versamlung, tab. 3.) Laeiniate 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 
Con [fen i', to such an extent as to give an appearance 
as if there were several cotyledons. 1 

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 in- 
stances 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 leafstalk 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 corre- 
sponding 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 

1 Duchartre, ' Ann. S<?. Xat.,' 3rd series, 1848. vol. x, p. 207. 




question. Many are cultivated as garden varieties 
under the names here given. 

Trollius europseus dissectus. 
Chelidonium majus laciniatum ! 
Glaucium luteum. 
Brassica oleracea ! 
Tilia parvifolia laciniata. 

asplenifolia ! 
Acer platanoides laciniatum. 

^Esculus Hippocastanum incisum ! 
Vitis vinifera apiifolia ! 
Ilex Aquifolium ! 
Rhus Toxicodendrnm quereifolium. 

Ervum Lens. 
Cytisus Laburnum quercifolium ! 

Rubus fruticosus laciniatus ! 
Pyrus communis. 
Cerasus Lauro-cerasus. 
Apium graveolens ! 
Pimpinella magna. 

Crataegus Oxyacantha laciniata. 

quercifolia ! 
Ribes nigrum. 
Sambucus nigra laciniata ! 

racemosa laciniata. 
Dipsacus fullonum. 
Scabiosa atropurpurea ! 
Sympboricarpus racemosus. 
Heliantbus sp. ! 

Lonicera Periclymenum querci- 
folia ! 
Syringa persica laciniata ! 
Syringa vulgaris ! 
Nerium Oleander ! 
Lamium purpureum. 

album ! 
Salvia officinalis. 

Solanum Dulcamara ! 
Fraxinus excelsior crispa. 
Veronica austriaca. 
Polemonium cseruleum. 
Juglans regia laciniata ! 

Anemiopsis californica. 
Chenopodium Quinoa. 
Ulmus americana incisa. 
Fagus sylvatica beterophylla ! 
laciniata ! 
aspleniifolia ! 
salicif olia ! 
Mercurialis perennis. 
Urtica dioica. 
Quercus Cerris laciniata ! 

pubescens filicina. 
Betula populifolia laciniata. 

alba dalecarlica. 
Alnus incana laciniata ! 

glutinosa laciniata ! 
Corylus Avellana heterophylla ! 
laciniata ! 
Carpinus Betulus incisa ! 

Castanea vesca heterophylla. 
Populus alba acerifolir 
Orchis sambucina. 
Arum maculatum. 
Filices sp. pi. 

See also Schlechtendal, 'Bot. Zeit.,' torn, 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 ;i normal occurrence as in 
the pel ;ils of mignonette (Reseda), or those of Ateme 
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 t divided 
petals of linlnjx arctums. Among other plants subject 
to this division of sepals or petals may be mentioned 
as having come within the writer's personal observa- 
tion, Ranunculus Lingua, R. acris, Pajxtcer somniferum, 
and others of this genus, Sa/ponaria sjp. DiantJms, 
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 

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 Up was divided into three segments per- 
fectly 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 

Fig. 29. — Flower of Oncidium sp. seen from the back, 
divided into three unequal segments. 

The lip is 

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. 1 

Moquin mentions having seen the stamens of Mat- 
thiola incana and Silene conica completely divided, 
each section bearing half an anther, exactly as happens 
in PolygalacecB. 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 fre- 
quently when those organs become more or less leafy, 
as in Trifolmm rejiens, and other plants to be hereafter 

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 

I Masters, ' Rep. Bot. Congress,' London, 1866, p. 136, tab. 7, f. 15, 16. 


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 charac- 
ters proper to the species under natural conditions, 
but lost by cultivation or otherwise. Thus, Mr. Buck- 
man accounts for " finger-and-toe" in root-crops on 
the principle of reversion to the wild form. 



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 1 in- 
cluded cases of this description under the category of 
" Disjunctions 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 primi- 
tively 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. 

1 Log. eU., p. 298. 


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 hyemaMs, 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 TrifoUum repens, Ranun- 
culus Ficaria, &c l 

Dialysis of the parts of the same whorl :— calyx. — The separa- 
tion 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 dia- 
grammatically by a dotted line above the letters 
denoting the sepals, petals, &c. When this change 
happens in the calyx we have the gamosepalous con- 
dition replaced by the polysepalous one, as thus 
represented : 

s s s s s 
instead of 

s s s s 

as in a calyx of five coherent sepals. 

Detachment of this kind occurs not unfrequently, as 
in Primula vulgaris, TrifoUum repens, &c. In Bosacece 
and Pomaceai this separation of the calyx is of the more 
moment, as it has reference to the structure of the 

1 Masters in Seemann's ' Journal of Botany,' 1867, p. 158. 


inferior ovary, as will be more fully mentioned here- 
after. Here, however, a ease recorded by M. J. E. 
Planchon may be alluded to 1 wherein a quince fruit (( '7- 
donia) was Burmountedby five leaves, the surface of the 
pome being marked by as many prominences, which 
apparently corresponded to the five stalks of the caly- 
cine leave-. In tins specimen, then, the inferior position 
of the ovary appeared to be not so much due to an expan- 
sion of the fruit stalk, as to the fusion of the hyper- 
trophied stalks of the sepals. Some of the malforma- 
tions 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 
Bosacecs, Pomacece, Umbelliferce, less commonly in 
Leguminosce, also in the following genera : — Primula, 
Symphytum, Gentiana, Gamparmla, &c. 

Dialysis of the corolla is likewise of frequent occur- 
rence, either partially or to such an extent as to render 
the corolla truly polypetalous. Among Labiake the 
upper lip of the corolla may be often met with partially 

Fig. 30. — Dialysis of the sepals and petals in Correct. 

1 • Bull. Soc. Bot. France.' t. xiii. 1S66, p. 234. 


cleft, as it is constantly in Phlomis biloba, or more 
markedly among the Lobeliacece. 

In the Composite, 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 

A more complete separation occurs not unfrequently 
in Campanula, Rhododendron, Phlox, Sfc. Figs. 30 and 
31 illustrate dialysis of the corolla; the first in Correa, 
the second in Campanula. 

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 
gamopctalous flower takes place most frequently. 


Verbasoum ! 

Campanula ! sp, 




Digitalis ! 

Phlox ! 


Coboea ! 


Rhododendron ! 


Erica ! 

Gentiana ! 



Azalea ! 

Primula ! 

Composite! sp. 


Lamium ! 

Lonicera ! 

Convallaria ! 

Convolvulus ! 

Lilium ! 


Colchicum ! 

Antirrhinum ! 

&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 Legwminosce and Gom- 
jjositce, as in Tragopogon, as related by Kirschleger, 
in Hypochaeris by Wigand, and in Coreopsis by Schlech- 

Dialysis of the carpels. — In the case of the carpels this 
disunion is more frequent than in the stamens. M. 
Seringe 1 figures carpels of Diplotaxis tenuifolia more or 
less completely separated one from the other ; indeed, 
this separation is very common amongst Gruciferce and 

Generally speaking, the disunion is complicated with 
frondescence — but not always so. I have, in my her- 
barium, specimens of Gonvallaria majalis, Gommelyna 
sp., and of Lilium auratum, in all of which the three 
carpels are completely disjoined, and present three 
1 ' Bull. Bot.,' pi. i, figs. 8—12. 


styles, three stigmas, &c, without any other change. 
Engelmann 1 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 im- 
perfect disjunction, the two bi-lobed carpels were 
changed into a nearly one-celled capsule ; 2 and 3rd, 
those cases in which the carpels are open and foliaceous. 
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 
Gerasus and Amygdalus }><-r.<icn ; nevertheless, fleshy 
fruits sometimes become disunited. I have seen a 
case similar to that mentioned by M. Alphonse de 
Candolle in 8olanwm esculentum, in which the pericarp 
became ruptured, and the placentas protruded. A 
like occurrence has also been observed in a species of 
Melastoma. 3 This is analogous to what happens in 
Caulophyllum and Slateria. Disjunction of the carpels 

Fig. 32. — Anomalous form of orange. 

' De Antliol.,' p. 37. 

Moquin, loc. cit., p. 305. 

' Neue Denkschr. der Allg. Schweiz. Gesell..* band v. pi. ii, p. 5. 


/ •> 

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 
angers to the fruit. Of one of these, Ferrari, 1 in the 
curious volume below cited, speaks thus : " Arbor pro- 
rusissima, quia dat utraque maim ; imo quia vere manus 
dat in poma conversis ; utque magis munifica sit poma 
ipsa convertit in manus." 

M. Duchartre 8 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 3 considers the rind of the orange as a pro- 
duction 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. 

Fig. 33. — Orange. Showing dis- 
junction of carpels, after Maout. 

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 

1 ' Hesperides.' auctore Ferrario, Rome, 16-16, fig. 415, pp. 213 and 
215. See also Michel, ' Trait e du Citronnier.' 

2 ' Ann. des Science Nat.,' 3rd series, 1844, vol. i, p. 294. 

3 ' Org. Veget.,' vol. ii. p. 41. 


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. 



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 con- 
genital, the word detachment would be an expressive 
one to apply to these cases, but as the change in ques- 
tion occurs quite as often from a want of union, an 
arrest or stasis of development, as from a bond 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 cor- 
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 " Disjunctions 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 j 

I c c c c c i 

would indicate the disjunction of the sepals from the 


carpels (c),in contradistinction to adhesion, which may 

be n'lii'csciitt'tl 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 circum- 
stances, that between the calyx and the ovary is per- 
haps the most common. The calyx adhcerens or superus 
is a structural characteristic to which all botanists 
attach considerable importance ; so that when excep- 
tional cases occur in which the calyx becomes detached 
from the ovary, becomes, that is, inferus or liber, a pro- 
portionate 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 Rosacea?, the change in question is very 
common, especially in conjunction with an elongation 
of the axis of the flower (apostasis) and with prolifi- 
cation, 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 Pomacece. 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 Oxyacomtha, in some blossoms 
of which the hollowed end of the peduncle still invests 

Fig. 35. — Proliferous Rose. Showing an absence of the visual dilatation 
of the flower-stalk, and other changes. 

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 ordi- 
nary 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 wood- 
cuts, figs. 36, 37, consisting of separate, superior ovaries, 
sometimes destitute of ovules, or, at other times, 
having two of these bodies. 1 

This condition accords precisely with the account 

' <Gard. Chron.,' 1865. p. 554; 1867, p. 599. 

SOI ITMN (»F i ALYX I l.'"M riSTIL. 


of the development of fche flowers in Pomacea as given 
by Payer, Oaspary, and others, so that the flowers 

Fig. 36. — Section through. Apple 
blossom, showing detachment of 
calyx from ovaries, absence of 
dilated flower-stalk, &c. 

Fig. 37. — Calyx detached from 
carpels in Apple. 

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 1 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 malforma- 
tions ; 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 

On the other hand, M. Planchon's specimen of the 
Quince before alluded to, not to mention other in- 
stances, 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 im- 
possible to say where the axial or receptacular portion 
ended, and the foliar portion began. As both from 

1 'Bull. Soc. Bot. Fr.,' 1854. p. 303. 


normal organogeny as well as from unnsnal confor- 
mation 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 Bosacece 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 
SaxifragacecB, Gucurbitacem, Myrtacece, BruniacecB, Rubi- 
acece, and other families of like conformation. 

In Umbelliferce, a detachment of the calyx from the 
ovaries frequently occurs, sometimes 
without any other change ; at other 
times attended by more serious altera- 
tions. 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 compo- 

Fig. 38.-Flower g j t j on f t h e fr^ 
of CEnantlie crocata, T tl -ni- tt7 7 7 • 

in which the five -I 11 the Kubiaceous genus nikkliict, 
sepals were com- as mentioned by Duchartre, the ovary 
iwt V tSlt is completely inferior, but when the 
here three in num- fruit arrives at maturity four small 
ber and destitute of i eaves are detached from its surface 
which had previously adhered to it, 
and which it seems reasonable to consider as the 

In CampcmulacecB a similar separation of calyx from 
the ovary maybe occasionally met with. On the other 
hand, the occasional formation of a leaf on the inferior 
ovary of these plants would indicate the axial nature 
of the fruit. In Gamjjanumcea and Gyclodon 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. panifiorum, placed on 
the peduncle of the flower far removed from the base 
of the corolla and ovary, whilst in G. truncation and in 

SOLI il"N OF ( AM \ FROM PJS1 ll.. -I 

./in ii a urn 'i they adhere fco the base of i Ik- tube of the 
corolla." 1 In this order, thru, as in SaxifragacecB, 
Bruniacece, &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 OnagracecB (Jus&ieua), as also in ( 'actacece (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 Epi- 
lobivm, 1 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 ' Xew Arrangement of Phanero- 
gamous Plants,' p. 4, cites a case wherein the perianth 
was completely detached from the surface of the ovary 
in GaimaMs 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 sub- 
stitution of carpels for stamens, as in Begonia, Fuchsia, 
&c. Among GucurbitacecB, examples have been re- 
corded, both of the detachment of the calyx from the 
ovary, 3 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. 

1 Hook et Thorns, ' Pra;ciirs. ad Flor. Lid.,' Journ. Linn. Soc. vol. ii, 
1858, p. 6. 

- Lindlcv. ' Veget. Kingd..' p. 315. 



The following are the genera in which the change 
has been most frequently observed : 

*Rosa ! Athamanta. 

*Pyrus! Selinum. 

*Oata3gus ! Carum. 

*Daucus ! Imperatoria. 

Pastinaca. Rudbeckia ! 

Torilis. ^Campanula ! 

Apium. Lonicera ! 

(Enanthe ! Cucumis ! 

Heracleum. 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 Gobaa scandens (Turpin), in Antirrhiwu/m 
majus, and in many double flowers. 

Partial detachment of the stamens from the styles 
occurs frequently in semi-double flowers of Orchis? 

1 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. Engehnann, 'De Antholysi,' p. 37, tab. v. C. 
Morren, ' Bull. Acad. Belg.,' six, part 3, p. 318. Cramer, 'Bildungs- 
abweichungen,' 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 Myrtacew), 
and other treatises on Organography. 

i'.\ EtT I i i 


lbili connected with changes in the arrange- 
ment 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 posi- 
tion of an organ ; while Heterotaxy may serve to in- 
clude 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 




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 tera- 
tological phenomena. It is obvious that abnormal 
unions or disunion-, 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 

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 carm/mosa? 

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 L&ueoivm cestivum, and the 
changes observed were apparently attributable to a 
simple separation of two leaves that are usually con- 
tiguous. " Suppose," >ays M. Gay in describing this 

1 'Tijdsclir. toot. nat. Gesdh.,' viii. 1841, tab. ii, p. 17- 



malformation, ° the first leaf of the terminal bud sepa- 
rated by a long internode from the other leaves, which 
remain closely packed; and further, suppose an evidenl 
thickening of the upper portion of the lengthened 
internode, and there will be not only a single bull), 

Fig. 39. — Unusual position of bulbs of tulip; the parent-bulb out open. 

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." 

The formation of bulbs in the axils of the leaves, as 
happens occasionally in tulips, is further alluded to 
under ihe 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 in- 
stances 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 spike- 
lets and the main stem ; in Triticum the spikelets are 
placed with their backs against the rachis, in Lolvum 
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 
Leucanthemivm pratense, in which the ligulate female 
flowers were growing singly in the axils of the upper 
leaves of the stem. 1 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 capi- 
tulum may often be observed in Composite, Dipsacece, 
&c. I have also met with specimens of Lamiv/m 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 

Caspary 2 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 

M. Founder mentions an instance in Pelargonium 
grandijlorum, 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 

1 Communication to the Intemat. Bnf. Congress, Paris, 1867. 
■ 'Bot. Zeit.,' 1859. p. 117. tab. v. 


axis. There were sever;]! bracta beneath this flower 
disposed spirally in the .. arrangement, all being empl v, 
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 some- 
times give rise to an apparent displacement, but the 

true nature of the malformation can, in general, be 
readily made out. 

Steinheil 1 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 dis- 
tinct 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 alter- 
nation. 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. 2 The term " diremption" has sometimes 
been applied to cases where leaves are thus apparently 
dragged out of position. 

1 ' Ann. Sc. Nat.,' ser. 2, vol. iv, 1835, p.. 1 13. tab. v. 
See Kirschleger, ' Flora,' 1844, p. 566 [Scabi 



In Tradescantia virginica I have met with opposite 
connate leaves ; the altered position, however, being 
due to the union of two stems. 

Twisting of the stem is a frequent cause of the dis- 
placement of leaves (see spiral torsion), as also hyper- 

Fio. 40. — Large-coloured leaf occupying the position of the infloroa- 
:enee it) Gesnera, after Morren, 



trophy, whether thai excess of development take place 
laterally or lengthwise (see elongation). Atrophy or 
suppression will also frequently bring about an altera- 
tion 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 sup- 
pression is that mentioned by Morren, 1 where, in 
Gesnera Geroltiana, a large leaf apparently occupied 

Fig. 41.— Ordinary arrangement of leaves in fascicles of three in 
Pinus pinea and unusnal arrangement of leaves of same plant in spires. 

1 • Bull. Acad. Belg./ t. xvii. part ii. p. 387. 


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, how- 
ever, 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 La/rim) or Pinjis 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 

Fig. 42. — Altered arrangement of leaves of yew, Taxus baccata. 

manner, the elongation of the shoot being arrested. 
This appears to be the result of the injury effected by 
some insect. 


So, too, the alteration from" verticillate to spiral, or 
vice versa, may take place without any other aotable 
change. 1 This may frequently be seen in Rhodo- 
dendron 3. 

Displacement of the parts of the flower. — Tins subject is 
partly touched on in the chapters on solution, ad- 
hesion, and in those on hypertrophy, elongation, 
prolification, &c, so that in this place it is only requi- 
site to oiler a few genera] remarks, and to refer to 
other sections for farther 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. 2 The same author 
also gives an account of the displacement of several of 
the organs of the flower in GypripediuTn 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 Orchidacece are by no 
means uncommon ; the following may be cited by way 
of illustration. In a specimen of Oncidiwm 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 dis- 
placement of the sepals and petals — so that all were 
dragged out of place. This dislocation may be better 
appreciate by the accompanying formula than even 

1 ' Clos. Mem. Acad. Toulouse,' 5th sev.. t. vi. pp. 51, 70. 
1 'Bull. Acad. Roy. Belg.,' xviii. part ii. p. 505. aud vol. xvii, part i, 
p. 190. aud vol. xix. part i. p. 260. 



by the woodcut. Let the usual arrangement be thus 
represented : 



s standing for sepal, p for petal, l for lip, ST for 
stamen ; then the dislocated form may be represented 
thus : 

s s 


Fig. 43. — Flowei" of Oncidium cucullatum, showing union of two lower 
sepals, displacement of column and lip, &c. 

In a specimen of Cyprvpedium 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 pecu- 
liar 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, bul the lateral petals are distorted, 
displaced, and adherent one to the other and to the 

Fig. 44.— Malformed flower of 

Fig. 45. — Diagram of mal- 
formed Cypripedium. o, outer 
segments ; i, inner segments of 
perianth ; e, lip ; g, stigma ; a, 

column, while the posterior shield-like rudimentary 
anther is completely wanting. 

In a specimen of Ly caste SMnneri similar changes 
were observed, as shown in the plan, fig. 47. Here 
the posterior sepal was deficient, the two lateral ones 

Fig. 46. — Diagram showing ordi- Fig. 47. — Plan of flower of Ly- 

nary arrangement in Cijpripedium. caste Skinneri, showing displace- 

o, outer, i. inner segments of peri- ment of organs, 
anth; e, lip ; a, anther; a', abortive 
stamen ; s. stigma. 

were present, one of them with a long tubular spur, oo ; 
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, /, one above and within the other. The 



column and ovary of this flower were in their normal 

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 

Fig. 48. — Plan of malformed 
flower of Dendrobiuvi nobile. 

Fig. 49. — Plan of natural arrange- 
ment in Dendrobium nobile. The x x 
represent processes of the column, 
perhaps rudiments of stamens. 

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 appre- 
ciated by comparing them with fig. 49, a diagram 
showing the natural arrangement of 
parts in this species. 

Sometimes the displacement seems 
consequent on hypertrophy of one of 
the parts of the flower, the dispro- 
portionate 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. 

It is curious to observe in many 
of these cases that the transposed organ not only 

Fig. 50.— Plan of 
flower of violet 
showing displace- 
ment of petals, &c. 
At b was a rudi- 
ment of a stamen. 


occupies the place of a suppressed or abortive organ, 
but frequently assumes its colour, and, to some extent, 
its function. 'Tin's 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 1 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 Shwmeri, thus recalling the 
structure of Masdevallia, where the labellum is normally 
very small. The arrangement in Lycaste may thus 
be symbolised : 

p st r 

s s 

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 guid- 
ing insects, is deficient, its place is supplied by other 

Displacement of the parts of the flower from elon- 
gation 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 Primmla sinensis and in the 
Pea [Pisv/m 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. 1 

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 dis- 

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 mar- 
ginal. Moreover, it frequently happens, when the 
carpels become foliaceous and their margins are discon- 
nected, 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 Rarwnculus Ficaria that 
came under the writer's notice the carpels were open, 
i. e. disunited at the margins, and each bore two im- 
perfect 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. 2 

The occurrence, also, of different forms of placen- 

1 See also Schlechtendal. ' Bot. Zeit..' iv. p. 804. Primula veris, parti- 
bus perigonii spiro: in modum conflucnfibus. 

' Seemann's ' Journal of Botany." vol. v. 1 R 67. p. 158. 



bation in differenl 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. 1 
Professor Babington describes in the ' Gardeners' 
Chronicle,' 1844, p. 557, a curious flower of Cerastium, 
in which, in addition to other changes, the five carpel- 
lary 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" 
(tig. 51). See also Lindley, ' Veg. Kingdom,' p. 497. 

Fig. 51. — 1. Monstrous flower of a Gerastvwm : sepals and petals 
leafy. 2. Stamens and pistils separate. 3. Ovary cut open to show the 
imperfect dissepiments and thp attachment of the ovules. 4. A deformed 

M. Baillon~ records flowers of Bunias, some with 
ovules on the margins of the carpels, others with a 
central branch bearing the ovules; hence he concludes 

1 ' Journ. Linn. Soc' i. 1857, p. 161, c. xylog. 
" Adansonia.' ii. 306. 


very justly that no fair inference can be drawn from 
these facts as to the normal placentation of Cruciferge. 

The same excellent observer has recorded the occur- 
rence of free central placentation in malformed flowers 
of Trifolium rep ens} 

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 Gesneracece. 

The same author alludes to certain chano-es 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 ex- 
planation of some cases of anomalous placentation, for 
the only indication of the true nature of the placen- 
tation 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 

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 pro- 
longation of the axis, and the formation on it of a 
second whorl of carpellary leaves. 

In double flowers of PrimulaceoB 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 placen- 
tation to another are not infrequent. The late Pro- 
fessor E. Forbes describes 2 an instance of true foliar 

1 ' Adansonia,' iv, p. 70, t. i. 

' Honfrey's • Bot. Gazette,' i, -JU.j. 


and true axile placentation in the same flower in I 

These and mam 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 

It will be remembered, also, how, in certain natural 
orders, under ordinary circumstances, considerable 
diversity in placentation exists, according as the mar- 
gins 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 Caryophyltece, Gucurbitcn 
Papav&racece, 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 

It is not necessary, in this place, to enter into the 
question whether the placenta is, in all cases what- 
soever, 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, how- 
ever, 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 in- 
ferences 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, ' Ami. Nat. Hist..* 1843, vol. xi, 35. Brongniart, 'Ann. Sc. 
Nat.,' 1834, ser. 2. i. p. 308. Alph. De Candolle. ' Neue Denkschrift dev 


Allg. Schweizer Gesellscb.,' Baud v, 1841, p. 9. Ducbartre, ' Ann. Sc. 
Nat.,' 3rd ser., 1844, vol. ii,p. 290. Ibid.. ' Elem. Bot.,' p. 574; 'Rev. 
Bot.,' 1846-7, p. 213. Babington, ' Gard. Cbron.,' 1844, p. 557. Lindley, 
' Elements,' p. 89 ; ' Veg. King.,' pp. 313, 497, &c. Berkeley, ' Gard. 
Cbron.,' 1850, p. 612. Unger, 'Nov. Act. Acad. Nat. Cur.,' 1850; 
and in Henfrey's, ' Bot. Gazette,' 1851, p. 70. Scbleiden, ' Prin- 
ciples,' English edit., p. 385. Payer, 'Elem. Bot.,' pp. 196, 211, 
224. Baillon, ' Adansonia,' iii, p. 310, tab. iv. Cramer, ' Bildungsab- 
weichungen,' p. 20, &c. Clos, ' Ann. Sc. Nat.,' 5tb ser., iii, 313, as well 
as any of tbe general treatises on botany. Reference may also be made 
to tbe cbapters on Prolification and Substitutions (in tbe case of tbe 
carpels and ovules), and to tbe authorities tberein cited. 



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 

The adventitious bud may be a leaf-bud or a flower- 
bud ; it may occupy the centre of a flower, thus termi- 
nating 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 sub- 
division 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 malfor- 


mation than the medial] form. If only the number of 
orders and genera be reckoned, the truth of tins state- 
ment will lie .scarcely recognised ; but if individual ca 
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 lias 
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 tlower-bud do not essentially differ. 1 
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 flow r ers 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 pro- 
lification, 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 floAver-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 espe- 
cially 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 

1 Linn,. ' Prolepsis,' § vii; Goethe, ' Metamovph.,' §§96, 103, 106. 


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 exclu- 
sion 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 inflores- 
cences. 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 in- 
definite ; 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 

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. 

rrolifieation of the inflorescence. — This consists in the for- 
mation 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 
organs, and from chloranthy, in which all or the grei 
portion of the parts of the Bower are replaced by lea 
Prolification is, in fact, a formation of supernumerary 
buds, leafy or floral, as the case may be, these bnds 
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 1 seeing that a similar condition is so 
commonly met with normally, as in Epacris, Metrosideros, 
JBromelia, Eucomis, &c, wherein the leafy axis projects 
beyond the inflorescence proper; or as in Primula vm- 
perialis, in which plant, as also in luxuriant forms of 
P. sinensis, tier after tier of flowers are placed in succes- 
sion above the primary umbel. Xevertheless, 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 GoniferoB, 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 irre- 
levant 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 Gryptomeria 
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 Gasta/nea 

■ Diaphysis Lnflorescentiarum," Bngelmann, ' De Anthol..' § 85. 


vesca, each of which had a tuft of small leaves at their 
extremity. In the common marigold and in Lotus 
cmmiculatus I have also seen instances of this kind. 
Kirschleger 1 describes a tuft of leaves as occurring on 
the apex of the flowering spike after the maturation of 
the fruit in Plant ago, 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 vir/i- 
■parum, in which not only did particular branches ter- 
minate in rosettes of leaves provided with roots, but 
similar growths proceeded from the heads of flowers 
themselves. Baron de Melicoq' 3 gives a case in 
Primula variabilis, in Avhich 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. 
\V. 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 Com- 
mittee of the Horticultural Society on July 11th, 1868, 
by Mr. Wilson Saunders ; the species in this case w T as 
P. cortusoides. To Mr. R. Dean I am indebted for a 
similar proliferous cyclamen, which seems similar to 
one mentioned by Schlechtendal. 3 This author alludes 
to an analogous circumstance in the inflorescence of 
Oytisus nigricans, where, however, the change was not 

1 ' Flora,' 1844, p. 565. 
" 'Ann. Sc. Nat.,' ser. 3. vol. v, 1846, p. 64, 
• B >t. Zeit.,' vol. xx, p. 382. 


so great as in the preceding cases. The instances just 

cited all occur in plants having an indefinite form of 

Fig. 52. — Inflorescence of Polyanthus, bearing a tuft of leaves at the 
top of the scape intermixed with the flowers. 

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 longi- 
tudinem adepti ftierunt." 1 

, 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 

1 ' Miscel. Austriac. Bot..' vol. i, Vindob. 1778. p. loo. 


wild celery and other Umbelliferce? 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 Eu- 
phorbia palustris." 

Lateral foliar proMcation of the inflorescence is of more com- 
mon occurrence than the preceding. I have met with 
it, amongst other plants, frequently in Brassica oleracea, 
Pelargonium zonule, Scabiosa, Bellis, and many other 
composites, also in Leguminosce, 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. 3 Many species of Allium, Zilium, Saxifraga, 
Begonia, AcMmenes, normally produce leaf-buds or bulbs 
in the inflorescence ; so, too, leafy shoots are sometimes 

i " Umbellatl dum proliftcantur, augend umbelhdam, ut ex umbellnla 
simplici altera exeat." ' Linn. Phil. Bot.,' § 124. 

2 ' En. Euphorb.,' p. 36. 

3 Meisner, ' Mon. Gen. Polygon i Prodrom.,' p. 20, tab. v, considers the 
bulbils of this plant to be modifications of the pedicels of the flower. 


round iii Mistmi natans, Juncus uligmosus, Chlorophytiwm 
Sternbergiarwm, &c. As an accidental occurrence, ;i 
Bimilar thing has been noticed in Lychnis corcmaria, 
Phai/us grandifolms, Oncidvu/m cebolleta, Epidend/rv/m 
elongatum? &c. &c. 

Here, loo, may be mentioned those cases wherein a 
leaf-bud is found upon the surface of the so-called in- 
ferior 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, 
i nude 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, Pereslcia, Crataegus banacetifolia, &c. 

The fruits of Opuntia Salmania and of 0. 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 Hete- 

Lateral floral prolification of the inflorescence. — This, which is 
termed by Engelmann Ecblastesis foliorum sub flora- 
lium, 2 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 

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 
Vmbelliferoi, Dipsaceo3, and Covipositce ; a familiar illus- 
tration in the latter order is afforded by the hen-and- 
chicken daisy. In some species of Composite, indeed, 

1 See A. Braun, ' Ann. Scienc. Nat.,' 4th series, 1860, vol. xiv, p. 13. 

- " Prolificatio e latere ex calyci communi proles plurimos pedunculatos 
LHiittens, fit in compositis aggregates proprie dictis." ' Linn. Phil. Bot..' 
§ 124. 



it is a normal and constant occurrence, while in other 
cases, such as Filago germanica, usually described as 

Fig. 53. — Lateral prolification in inflorescence of Pelargonium. 

proliferous, there is not, strictly speaking, any prolifi- 
cation, 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. Schlech- 
tendal 1 gives a summary of the various kinds of malfor- 
mation affecting the inflorescence in Plantago, and 
divides them into five groups, as follows: — lst,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 

1 ' Bot. Zeit,,' 1857, p. 873. See also « Yerhandl. Nat, Hist. Vereins. 
Freaks. Rheinl. u. Westphal./ 1854, t. ix. 



head with other rosettes. 5th, paniculate, in which the 
inflorescence lias become a much-branched pyramidal 
panicle, covered with little bracts, and with very rudi- 
mentary flowers. 1 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 bor- 
ders of richly cultivated fields in the 
neighbourhood of London I have fre- 
quently gathered specimens of Plantago 
major with a branched spikeprovided 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 Schlechtendars 
bracteate and polystachyate divisions. 
Wigand 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 sec- 
tion, 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 

Fig. 54.— Plan- 
tago major, with 
panicled inflo- 

1 " Pannicula spicatim spaxsa onusta immrnera fcetura herbaeeorum 
flosculorum racematim coha?rentium," ' 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. 

• ' Flora.' 1856, p. 706. 



watching the growth of the rose plantain must have 
observed the occasional production of flowers, sometimes 

Fig. 55. — Inflorescence of Plantago major, with bracts partly replaced 
by leaves and spike branched. 

stalked, in the axils of the leafy bracts, and at the same 
time have noticed that the internocles become elon- 
gated, 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, 1 wherein were several 
spikes, some sessile, others stalked and pendent, the 
whole intermixed witli 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). 

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 PI aula go 
seems to have its own perverse mode of growth ; for 
instance, the bracteate, polystaohyate 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 pro- 

1 • Flora of Berwick-on-Tweed,' vol. i. p. 38. 



lification of the inflorescence. As, however, there is 
still some difference of opinion as to the correct mor- 
phological 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. 

Some of the cases of prolification of the inflorescence 
resulting in a branching of an ordinarily simple inflo- 
rescence, as in Reseda luteola (fig. 57), might equally 
well be placed with fission or multiplication of the 

Fig. 57.— Branched inflorescence of ifoWo hdeoln. 

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, 1800, p. 121, and the same author gives an 
illustration in his ' Orchidographia Europrea,' tab. 1 •"><>. 

In Grasses, as indeed in other plants with aspicate 
inflorescence, this change occurs not unfrequently. 
The common Ray Grass (Loluim) is especially subject 
to the change in question, and among cultivated 
cereals, maize and wheat occasionally show this ten- 
dency 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 fre- 
quently observed in the following genera : 





Ranunculus ! 




*Brassica ! 


Lychnis ! 
Diantlrus ! 


*Pelargonium ! 



*Trifolium '. 

Trifolium '. 

Lotus ! 

Lotus ! 

Coronilla ! 





*Pyrus ! 

*Pyrus ! 

*Cratsegus ! 

Crataegus ! 

*Rosa . 

Rosa ! 








? Tetragonia. 





Saxifraga ! 

! T m belli ferre. 


*Apium ! 



Heracleum ! 

Heracleum ! 











Angelica ! 






Begonia ! 




*Scabiosa ! 

*Scabiosa ! 

Knautia ! 

Knautia ! 


*Bellis ! 









Tragopogon ! 

Tragopogon ! 
Rudbeckia ! 
Senecio ! 

Bidens ! 











Azalea ! 


Convolvulus ! 

Convolvulus ! 
Calystegia ! 

Scr ophulariaceae . 

Scropbularia ! 
Antirrkiniim ! 


Acliimenes ! 


Priinula ! 

Primula ! 

Cyclamen ! 

Cyclamen ! 



Plantaginacese . 

*Plantago ! 

*Plantago ! 


Polygonum ! 


Eupliorbia ! 




Corylus ! 

Castanea ! 



*Larix ! 
*Cryptomeria ! 

Taxodium ! 



Phaius ! 
Epidendrum ! 
Oncidium ! 

Opbrys ! 


*Allium ! 
*Ornitliogalum ! 





*Lilinm ! 




Alisma ' 


*Jun< u ! 


Restio ! 

Restio ! 

Elegia ! 

Elegia : 

Willdenovia ! 

Willdenovia ! 




*Lolium ! 
*Zea : 
*Triticum ! 
*Hordeiim ! 

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. Engelinarm, ' De Antliolysi,' 
§§ 85—87. Fleischer, ' Missbild. Versch. Cultur. Pflanz.' For figures 
ot'Hen and Chicken Daisy (Bellis prolifera), see Lobel, ' Ic.,' 477. Sweert. 
' Florileg.,' pi. 98, f . §. ' Hort, Eystett. Plant. Tern.,' 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.,' pi. aestiv. fol. iii, f. i. Klinsmann, ' Linnsea.' 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. Sic. ' Hort. Eystett.,' pi. eestiv., t. vii, f. 2, P. 
rosea etP. bracteata. Lobel, ' Stirp. Hist.,' p. 162. Dodonseus, ' Pempt.,' 
1 — t, cap. xxiii, P. major spica multiplex, *. e. paniculata. Gerard, 
'Herbal.' Clusius, 'Plant. Rar. Hist.,' lib. v, p. 109—10, Plantago 
augustifolia Gar eti prolifera. Marchand, ' Adansonia,' iv, p. 156. 

Coniferce. — 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. fern, struct, morphol.' Parlatore, ' Ann. Sc. Nat.,' 
1862, vol. xvi, p. 215. Cramer, ' Bildungsabweich.,' p. 4, &c., &c. 

Gramineoj. — 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 con- 
stituent 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), represent- 
ing the thalamus of a strawberry 
prolonged beyond the fruits into 
a small leaf-bearing branch. 

Fig. 58. — Receptacle of straw- 
berry prolonged into a leafy 
branch. From the ' American 
Agriculturist. ' 

Fig. 59. — Flower of Verbascum, 
with five disunited sepals, five 
similar green petals, and a pro- 
longed 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 
bo Mi 1 . Salter, t here was a gradual change from the 
floral to the foliar condition; thus there were five dis- 
tinct lanceolate sepal-, 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 1 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. 

1 " Adansonia,' vol. iv. 1864. p. ISO, tab. vii. 'Gard. Chron.," 
November 19th. 1864. 


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 

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 approxi- 
mated, the ovary was open above, and sent out six 
shoots from within, perfectly developed, clearly repre- 
senting the central placenta and five axile buds, and 
each p'ivine out a number of adventitious roots at its 

In other genera of the same order (Primula our) an 

ILIPIOATIOB 01 Nil: J LOWBB. 1 1'.' 

tension of the placenta into a leafy branch has been 
observed, as in Lysima 7 via t where in one case the pro- 
longed placenta was removal and struck as a cutting. 1 

In Ericacecs too, the axile placenta has been 
©villiferous at the base and prolonged above into a 
leafy branch." 

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 Bamunculus affected with me- 
dian floral prolification. 3 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 calvx, 
corolla, and stamens, but with no trace of pistil. I 

1 ' Ann. Sc. Nat.,' ser. 3. torn, ii, p. 290 ; and ' Adansonia,' iii, tab. iv; see 
also Bureau, in ' Bull. Soc. Bot. France.' x, p. 191. 

- Baillon, 'Adansonia.' i. 2^0. 

3 See also figure in ' Hort. Eystett. Ic. Plant. Vera.,' fol. 15, fig. 1. 
Ranunculus asudicus, 



have been unable to find recorded any instance of mal- 
formation among Labiates or Borages at all similar to 

Fig. 60. — Flower of Diuntfms affected with median floral prolification. 

this. It differed from most other examples of prolifi- 
cation in that the axis was not prolonged, the adven- 
titious 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. 

In a flower of Conostephium (Epacridacece) 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: Ranunculacece, Garyo- 


phyllacece, Rosacea; while ii is commonly mei with in 
Scrophulariaceas, Primulacece, and I r mbelliferce. ( )f genera 
which seem peculiarly liable to it may be mentioned 
the following: Anemone, Ranunculus, Gheiranth/us, 
Dianthus, Dictamnus, Daucus, Rosa, Oeum, Pyrus, 
Trifolium, Antirrhvrmm, Digitalis, Prwmla. 

A reference to the subjoined list of genera affected by 
this malformation, and theknowledge 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 
wry 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 inflo- 
rescence, 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 PrimulacecB 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 ; l in Thesium also, another genus with 

1 Duchartre, ' Ann. des sc. nat..' 3me serie, vol. ii. 1844. p. 29o. 


free central placenta, this malformation has been 

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 Umbelli- 
ferre, where the carpophore may be truly considered an 
axile production; it is common among Bosacece and 
Ranuneidacece, 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 Magnoliacece, 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 termi- 
nated by a flower ; it is noticeable, also, in these speci- 
mens, 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 Leguminosce and in Santalacece. In 
Leguminosce, as also in Amy g dolus, it would seem as if 


tlio adventitious bud were strictly a lateral and axillary 
production, and moreover that the carpel itself is not 
Btrictly terminal but lateral in position, though appa- 
rently terminal from the abortion of other carpels. In 
the only recorded instance that I am aware of, of this 
malformation affecting the genus Thesvum, the pistil 
was altogether absent, and occupying its place was 
the new bud or branch. 1 

Fig. 61. — Daucus Carota, showing leafly carpels, prolification, &c. 

1 Reissek, ' Linrtsea,'* vol. xvii, 1843, p. 641, tab. xix. 


As the carpels are not imfrequently absent in cases 
of median prolification, it Las been thought that the 
pistil in such cases was metamorphosed into a stem 
bearing leaves or flower.-. Setting aside the physio- 
logical 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 un- 
altered 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 Da/ucus Ga/rota gathered in Switzer- 
land (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, sur- 
rounded by minute bracts. 1 

1 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 : — 

111 111 

3 2 3 2 3 3 2 3 2 3 

11111 11111 

The lower line of figures represents the calycine ribs, the middle row 


Not only arc 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 Dan ens 
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 prolifi- 
cation of the flower of AncJmsa ochroleuca, in which 
the pistil consisted of two leaves, situated antero-pos- 
teriorly on a long internode, with a small terminal 
flower-bud between them; and numerous similar in- 
stances might be cited. 

In this place may also be noticed those instances 
wherein the placenta elongates so much that the peri- 
carp 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 Solarium have been put on 
record by M. Alph. de Candolle. 1 This is a change 
analogous with that which occurs in some species of 
Leontice or Ccwlophyllum, 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, 

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 men- 
tioned, but have no direct upward prolongation into the carpel. The 
ten ridges are placed opposite to the sepals and petals. 

1 ' Neue Denkschriften der allgemeine Schweizerischen Gesellschaft,' 
bandS. 1841. tab. 2. 



become carried up with the prolonged axis, more widely- 
separated one from the other than below, and particu- 
larly liable to undergo various petalloid or foliaceous 
changes as in proliferous Roses, Potmtilla, &c. 

Fig. 62, copied from Cramer, shows an instance of 

Fig. 62.— Median floral prolification, <fcc, in flower of Delphinium. 

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 

PROl.ll'C ATI"\ OF THE FLOW1.I. L2< 

uxied 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 
1 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 UmbeUifercB. In a species of Gamparmla 
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 enclos- 
ing 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 Camparmla, 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 Composite 
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 
IiudbecMa? 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 in- 
creased 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 aloug 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 Orchidaceoe, 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 

Fig. 63. — Median prolification in Orchis pyramidalis, the outer seg- 
ments of the perianth regular and reflexed. 

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 

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 Diyitalix pur/tinra; 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 occu- 
pied 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 com- 
paratively rarely altered ; for instance, in proliferous 
pinks (Dianthns) the calyx is seldom affected, except, 
indeed, in those instances where the floral axis is pro- 
longed, 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 imper- 
fect 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. 

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 elon- 

PROLIFICATION OF THE l'l.uv> ] i) | 

gated (fig. 64) after the fashion of G&u/m rivale, &c.' 
Occasionally from the middle of the outer surface of the 
urn-shaped thalamus proceeds a perfecl 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-. L2. 

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 pro- 
liferous flowers of Umbelliferce s Garn/parmlacece, Corn- 
ice, &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 ; 9 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 sta- 
mens and pistils, according to the particular plant 
affected. Occasionally in prolified flowers the parts 
of the corolla, like those of the calyx, become folia- 
ceous, and in the case of proliferous pears fleshy and 
succulent. There is in cultivation a kind of Chevran- 
thus? 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 

1 Bell Salter. ' Gard. Ckron.,' March 13th. 1847, and ' Ann. Nat. Hist.,' 
1847, vol. xix, p. 471. &c. 

- ' The Origin and Production of Proliferous Flowers, with the Cul- 
ture at large for raising Double Flowers from Single, and Proliferous 
from the Double.' By J. Hill. M.D. London, 1759. 


ten or a dozen of these imperfect flowers may be 
seen on the end of a flower-stalk, giving an ap- 
pearance 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 

The stamens are subject to various changes in pro- 
lined 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 
UmbellifercB 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 sepa- 
rate 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 con- 
fined to them, as it is recorded among Boraginece. 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 PrimulacecB and other orders with 

free central placentation. 

Dnchartre, in his memoir on the organogeny of 
plants with a free central placenta, in the 'Ann. des 
Sc. Nat.,' 3 ser., 1814, p. 2 ( .><> 5 among other similar 
instances, mentions two flowers of Cortusa Matthioli, 
wherein the placenta was o villiferous 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. 1 In some flowers 
of Rhododendron I have observed a similar condition 
of the ovules, which, moreover, in the primary flower.-, 
were attached to the walls of the carpels — parietal 

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 pro- 
trude beyoud the carpels, or at least very rarely. 
There are, however, numerous instances of such an 
extension of the placenta and of prolification occurring 
among Primulacece in conjunction with the more or 
less complete arrest of growth of the carpels. 3 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 

1 A. de Candolle, ; None Denkschriften,' op. cit, p. 0, also Unger as 
cited in 'Botanical Gazette.' May, 1351. p. 70. 
: Duchartre, op. cit. 


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 prolincation 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 re- 
peated, 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 prolincation in no wise 
differs from ordinary central prolincation. Some- 
times 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. 1 In other cases, 
doubtless, the carpellary whorl is alone repeated, the 
other whorls of the adventitious flower being com- 
pletely absent. 

Another condition, apparently sometimes mistaken 
for prolincation of the fruit, is that in which the car- 
pellary 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 prolincation. This 
happens frequently in oranges, as in the variety called 
Mellarose. 2 

1 • Ann. So. Nat.,' 1844, vol. i. p. 297. 

- Maout, 'Lecons Elementaires de Botanique,' vol. ii. p. 48f> : Ferrari, 
• Hesperides,' pis. 271, 315, 405, 

PBOLIFIl a 1 1 - ■ N 0] I'ili: I BUIT. 135 

Moquin has given an explanation of the St. Valery 
Apple-, wherein the petals arc sepaloid, the Btamene 
absent, and where there is a double row of carpels, 
by supposing tlie<e peculiarities to be due to "a 
prolification combined with penetration and fusion 
of two or more flowers," but it is surely more rea- 
sonable 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. 1 

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 pro- 
ducing leaves. In these cases there is no true pro- 
lincation, 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 

1 Moquin-Tandon, loc. cit., p. 386, &c. ; see also Trecul, in the 'Bull. 
Soc. Bot. France,' torn, i, p. 307. 


before mentioned. Moqnin, 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 pro- 
lification 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. 

Ranunculacese. Clematis. 

Anemone ! *Anemone ! 

Ranunculus ! *Ranunculus ! 



Aquilegia ! 
Crucifera?. Bunias. 

*Cheiranthus ! 


*Matthiola ! 

Sisymbrium ! 

Brassiea ! 



Sinapis ! 






'I,u\\ BE. 



Cardaminc ! 


Heliantbemum ! 


Dianthua ! 

*Diantbus ! 
Silcne ! 
Lycbnis ! 


Viola ! 


Triu mfetta ! 


Geranium ! 


Pavia ! 

Pavia ! 


Hibiscus ! 


Byrsonima ! 


Genera not specified. 

*Dictamnus ! 


Caylussa ! 


*Citrus ! 









*Daucus ! 



*Rosa ! 

*Rosa ! 

*Geum ! 

*Geum ! 


Primus ! 


Spiraea ! 

*pyrus ! 

*Pyrus ! 

? Leguminosaj 

Trifolium ! 
Medicago ! 
Pisum ! 








Epilobium ! 


Epacris ! 


Rbododendron ! 









Jasminum ! 

Scropbulariacea? . 

Verbascum ! 

Antirrbinum ! 
*Digitalis ! 
*Linaria ! 















Genera not specified. 

* Anagrams ! 

Hieracium ! 



Genera not specified. 

Genera not specified. 



Phlomis ! 





*Anagallis ! 



Hieracium ! 


Calendula ! 




-*Campanula ! 



Tulipa ! 

Hemerocallis ! 


Hyacinthus ! 


Narcissus ! 


Orchis ! 




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 occur- 
ring within the flower from a similar state origi- 
nating 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 t be flower, or from the inflorescence beneath 
the flower. 

The accessory bud presents itself as a leaf-bud, a 
biancli, a iiower-bud, or a miniature inflorescence; 
it may be sessile, but is far more frequently stalked, 
and in more than half tlie number of cases it is a flower- 
bi id 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 adventi- 
tious growth occurs most frequently are the following : 
— Oruciferce, especially the genus Brassica ; Garyo- 
phyllacecB, e.g. Dianth/us ; Resedacece; Legwnmosce, 
e.g. Melilotus, Trifolium, &c. ; Rosacece, e.g. Rosa, 
Potentilla, &c. ; Umhelliferai, and Campcmulacece. 
For the most part, these are groups also peculiarly 
liable to central prolifi cation. 

All the parts of the flower may be thus affected ; 
but, as .might have been anticipated from the folia- 
ceous 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 dis- 
united 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. Gene- 
rally, however, the position of the accessory bud is 
such that it may property be referred to the axil of 
an undeveloped or rudimentary stamen. 

The largest number of instances of this malforma- 
tion, 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 relation- 
ship, 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 inflo- 
rescence 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 inflo- 
rescence. If individual instances could be accu- 
rately computed, the proportion would be even 

Fully three fourths of the entire number of genera 
recorded as occasionally the subjects of this irregu- 
larity 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. Garyophyllacece, Potentilla, Anemone, 
Bidamnus, UmbellifercB, &c. About one fourth of 
the genera have numerous stamens or numerous 
carpels, or both, springing naturally from the thala- 
mus. In others (about one sixth) the thalamus is 
enlarged into a diso, or else presents one or more 


glandular Bwellings, e.g. Reseda, Nywvphcea, QrucifercB. 

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 outer- 
most 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 propor- 
tion 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 occur- 
rence has been noticed in Galtha palustris, Herreria 
parvijiora, and other plants. Dr. Marchand's flowers 
of AnagaUis, previously referred to at p. 117, showed 
good illustrations of this occurrence, as also some 
specimens described by Kirschleger in A. phcenicea. 1 
Steinheil has figured and described 2 a flower of Scabiosa 

1 ' Bull. Soc. Bot. Ft.,' 1863, vol. x, p. 461. 

2 ' Ann. So. Nat.,' 1835, p. 65. See also Le Maout, 'Lecons Element.,' 
vol. ii, p. 4*26. 


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 Rosacea?., Pomacece, 
Philadelphia , or Tetragonia expansa, as elsewhere men- 

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 1 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 morpho- 
logical 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 
Gruciferce 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 

1 • Proc. Hort. Soc.,' vol. i, p. 39. fig. 2. 

PROL1FI0ATIOH 01 THE n.<»\\ ER. 1 fc3 

with the petals, which, together with the stamens and 
pistil, remained unaffected. The Dumber of parts in 
i lir supernumerary structures is generally Less than 
the normal flowers. 

In Mr. Eerbert Spencer's 'Principles of Biology,' 
part iv, p. :»7, are figured and described some mon- 
strous inflorescences in Angelica and other Umbelliferce, 
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. 1 

In the ' Gardeners' Chronicle,' 1855, p. 551, an 
instance is figured of the production of a supernume- 
rary flower proceeding from the axil of a stamen in a 
species of Nym/phcea (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 Nymphcea Lotus, or 
a cultivated variety of it. 

M. Wesmael 2 describes a very singular case of what 
appears to have been referable to axillary prolification 
in the flowers of Car ex acuta. The rachillus is de- 
scribed as prolonged through the utricle by the side of 
the stigmas, bearing on its side a bract, then a secon- 
dary utricle, from the axil of which sprung a short stem 
surmounted by an ovary. Wigand, ' Flora,' 1856, 
mentions a similar change in Garex 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 

1 See also ' Nat. Hist, Review,' 1865, p. 377. 

2 'Acad Roy. Belg.,' April 11th. 1863. 



female below and male above, others, again, wholly 

Various changes in the form and arrangement of 

Fig. 65. — Flower of Nymphcea Lotus, var., showing axillary floral pro- 
lification. The section also shows the tuft of leaves that occupied the 
place of the ovary. 

the several floral whorls accompany axillary prolifica- 
tion ; 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 modi- 
fication. The changes most commonly met with are 
such as may be classed under Goethe's theory of re- 
trograde metamorphosis ; for instance, if a supplemen- 
tary bud be developed in the axil of a sepal, that sepal 
is likely to be more than ordinarily leaf-like in appear- 
ance. The dislocation of the affected sepal from its 
fellows is a very frequent occurrence ; in cases of this 

PROLIPICATIOK OF THE ri.<>\\ i.i;. | |."> 

kind iho detached sepal ia placed below tin- others, 
thus approximating, in position as well as in function, 
to the bracts. In Borne of the instances of proliferous 
pears, on which 1 shall have occasion to comment, the 
sepals are described as sharing in the succulent cha- 
racter of the fruit. 

The petals, under such circumstances, often exist 
in the guise of sepals or of small leaves; and instanc» - 
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 
Trifoliwm 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 
si aniens, and in the centre a pistil in the guise of a 
trifoliate leaf. 1 Such instances seem to afford an ex- 
treme degree of a more common change, viz., the 
diminished size and contracted appearance of the 
sepals and petals when affected with axillary prolifica- 
tion. They have also a close relationship to such 
developments as we see in the wheat-ear carnation, 
in certain species of the genus Mcesa 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 compen- 

So far as the andraecium is concerned, the stamens 
either remain unaltered, or they arc 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. 
C6, where the stamens were entirely absent, and their 
places supplied by flower-bearing branches. This 
DiantJms has the more interest from its similarity to 
the one described by Goethe, Metam. der Pflanzen. 

1 ' Bull, Soc. Bot. Fr.,' torn. iii. 1S5C, p. 4-79. 




cap. 16, sect. 105 ; but in that instance median pro- 
lificatioii also existed. For my specimens I am in- 
debted to Mr. T. Moore. 

Fig. 66. — Flower of Dianthns 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 func- 

It occasionally happens that an adventitious bud 
arises from the axil of a monocarpellary pistil. This 
takes place sometimes in LegwmmoscB, and seems to 
have been more frequently met with in Trifoliv/m 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 ha* been placed in the axil of a 

PBOLI] [l LTION "I 1 l Hi: N. "W BE. I I , 

more or less Leaf-like carpel; while at other time 
second imperfecl carpel has been met with in the axil 
of the first. 1 I have myself seen numerous imperfectly 

(lt'\clc|ii'(l cases of t liis kind. 

It may be asked whether such cases are not more 
properlv referable to central prolification — whether the 
axis is not in such flowers terminated by two, rather 
t han by one carpel ? It is, however, generally admitted 
by morphologists that the solitary carpel of Legummosce 
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 in- 
stances of the presence of two and even of five carpels 
have been described. - 

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 3 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 prolonga- 
tion 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 

' 'Linnsea,' vol. xv, p. 266, c. ic. Caspary, 'Schriften d. Physik.-Oek. 
Gesell. zu Konigsberg,' bd. ii, p. 5, tab. iii, fig. 39, &e. 

2 Lindley, ' Veg. King.,' p. 545 ; also Clarke on the Position of* 
Carpels, Linn. Soc.,' December, 1850. 'Proc. Linn. Soc.,' ii, p. 105. 

3 ' Notuke,' vol. i, Dicot. p. 127. 'Atlas.' pi. xliii. 






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Complicated prolification. — From wliat 1i;is been before 

itated it may be seen that prolification of two or more 

kinds may coexisl in the Bame flower. Mixed leafy 

and floral prolification is not unfre<|iient in proliferous 

Fig. 67. — Proliferous Rose. Calyx leafy; petals normal, some re- 
flexed ; stamens and pistil absent ; in their places a branch with leaves 
and flowers. 

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, desti- 
tute, indeed, of calyx, but provided with numerous 
petals, stamens, and pistils. Above these lateral 

Fig. G8. — Rose exhibiting median, axillary, lateral, floral, and leafy 
prolification in same flower. 

flowers, the prolonged axis bore a number of scales in 
many rows. The scales were in their turn surmounted 



by ;i whorl of five perfect leaves, beyond which, again, 
the axis was prolonged into a leafy shool terminated 
by a flower bud, the whole constituting a remarkably 

complicated admixture of elements belonging to the 
flower, the bud, the inflorescence, and the leaf- 
shoot. 1 

Proliferous flowers of Orchids also occasionally pre- 
sent 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 follow- 
ing summary is extracted : 

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 



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 

1 Moquin-Tandon gives the following references to cases of proliferous 
roses, but some I have not been able to verify. 'Joum. 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. 



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 prolifica- 
tion of both kinds, median and axillary. The tertiary 
median growths, like the primary flower, did not 
develop median buds, but only lateral ones — quater- 
nary 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 

Fig. 70. — Diagram to explain the construction of the double-flowered 

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 

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'. 

In Narcissus major a similar combination of both 
forms of prolification exists, as described by Morren. 1 

On the general subject of Prolification in flowers, in 

1 ' 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. 


addition to the authorities already cited, the reader 
may refer bo the following among many others : 

[jinneeus, ' Prolepsis,' §§ \i el \ii. Goethe, 'Versuch. Metamorph.,' 
cap. svandxvi, §5 103—106. Moquin-Tandon, • El. Ter. Veg.,' p. 362, 
&c. Elngelmann, ' I).' Antholys.,' §§ 52—62, &c. Cramer, 'Bildungsab- 
weichungen,' &c. Orchidacece, UmbeUiferce, Compositce, Leguminosw, 
Primvlacet . Ranunculacece. Fleischer, 'Missbild. Cultur Gewachs.' 
Schlechtendal, ' Linnaea, 5 xv, p. 408, Rosa. ' Bot. Zeit.,' vol. xx, 1862, 
p. 882, Cyclamen. 'Bot. Zeit.,' vol. xx. p. 301, AspJiodelus ; etLilivm. 
Seringe, 'Bull. Bot,.' i. t. xi, f. 7, 8, Arabis, Diplotaxis. Clos, 'Mem. 
Acid. Toulouse,' 5th ser., 1862. Papaver. "Wigand, 'Flora,' 1856, p. 
TIC. HypocJueris; et ' Bot. Untersuch.,' p. 19. Buehenau, 'Flora,' 1857, 
p. 295, Reseda. Roeper, 'Bot. Zeit.,' 1852. p. 427, Orchis. Presl., 
' Linnsea,' 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. Klinsinann, ' Linnsea,' x, p. 604, t. v, 
Hesjperis. Fnckel, 'Flora,' 1848, p. 609, Melilotus. De Candolle, ' Or- 
ganogr.,' 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, Diunthus, 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 Euphor- 
biacearum et Capparidearum." Duhamel, ' Phys. Arbres.,' Hv. iii, cap. 
3, p. 303, pi. xii. i. 306, Rosa. Caspary, ' Bull. Soc. Bot. Ft.,' 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, Hyacinth us. Duchartre, 'Bull. Soc. Bot. France,' 1861, p. 451, 
Rosa. Weber, ' Verhandl. Nat. Hist. Verein. Rhein. Preuss., &c.' 1858 
et 1860. Landiin, ' Mem. Soc. Sc. Nat. Seine et Oise,' 1866 ? l 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 

1 I have not been able to meet with this, but it is said to contain a 
paper on prolification, with numerous bibliographical references. 


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. 1 

For other cases of prolification or the adventitious 
formation of buds on leaves, roots, &c, see under 



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 circum- 
stances, they would not be formed. It thus does not 
include cases of substitution, where one part is re- 
placed 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 

' • El. Ter. Veg.,' p. 364, Adnot. 

\i>\ i:\ii pioi s i,.,,, 157 

considered bo conic within the scope of teratology, or, 
;it 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 ordi- 
nary 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 con- 
sidered 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 

" Qidn et, caudicibus seclis, mirabile dictv, 
Trudiiur e sicco radix oleaghui 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 forma- 
tion 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 struc- 
ture 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 com- 
pensate 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. 

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 

Fig. 71. — Production of adventitious roots from leaf stalk of celery. 

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, 1 

i ' Gard. Chron.,' 1852, p 5] 

\h\ K\ CITI01 S BOOTS. 


and also in a cabbage whore a snail "having gnawed 
.i hole into the middle of a leaf at its junction with 
fche stein, 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. " 

The production of adventitious roots is not limited 
to the ordinary leaves of the plant, but may be mani- 

Fig. 72. — Germinating plant of mango, showing production of roots 
from one of tlie cotyledons (from the Kew Museum). 

fested on the cotyledons ; thus Irmisch describes cases 
of this kind in the cotyledons of Bunium cretimm and 
Carum Bulhocastanum. 1 I have figured and described an 
analogous case in the cotyledons of the Mango (fig. 72)." 
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 

1 ■ Flora,' 1858, pp. 32-42. 

2 ' Journ. Linn. Soc' vol. vi ; " Botany." 1862, p. 24. 



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. 

Layering is another garden 
operation dependent on the 
formation of these organs, and 

Figs. 73 and 74 show formation of roots from leaves induced by the 
art of the gardener. 

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 in- 
ducing 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 charac- 
teristics 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 Ja/ponica, Anemone Ja/ponica, &c. What are 
termed suckers, owe their origin to buds formed in this 

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. 1 From 
fche 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 

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" bear- 
ing the cotyledons, while at other times it is of con- 
siderable size. Generally it does not give origin to 
shoots or leaves other than the seed-leaves, but occa- 
sionally shoots may be seen projecting from it below 
the level of the cotyledons. This happens frequently 
in seedling plants of Anagallis arvensis, Euphorbia 

1 ' Tijdschvift vooov Natnm\ Geschied,' 1S3G, vol. iii, tab. vii, p. 171. 




p&plus, and other species, Tjmaria vulgaris, some TJm- 
belliferce, &C. 1 

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 extra- 
ordinary 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 pro- 
ceeding from the surface of the ovary or fruit, as in 

Fig. 75. — Leaf proceeding from 
hip of the Rose. 

Fig. 76. — Leaves proceeding 
from the ovary of Nymphcea. 

Crataegus tanacetifolia, roses, pears, gooseberries, &c. 
In a specimen of Nyw/phcea alba I have met with scale- 
like leaves projecting from the surface of the fruit (or 
torus?), and which did not appear to be metamor- 
phosed stamens or styles (fig. 76). 

1 Rceper, 'Enum. Euphorb.,' p. 19. Bemhardi, ' Linnaea,' vii, p. 561, 
tab. xiv, f . 1. Wydler, " Subcotyled. sprossbildung," ' Flora,' 1850, p. 
337. Hooker. 'Trans. Linn. Soc.,' vol. xxiv, p. 20 Welwitschia . 

\h\ i:\Ti I'Imi s LEAVES. 

For other illustrations of increased Leaf-formation, 
see Multiplication of foliar organs. 

Production of leaves on a usually leafless inflorescence. — The 
development of the bracts of an inflorescence to such 
an extent thai they resemble ordinary leaves is else- 
where alluded to as of common occurrence. It happens 
far less frequently that leaves are developed on an in- 
florescence usually destitute of them, without any 
metamorphosis or substitution, and without any forma- 

Fig. 77. — Leontodon. Scape with two leaves; the bracts of the invo- 
lucre are also leafy. 


tion of adventitious buds, such as happens in prolifi- 
cation. Such a partial change from a floriferous to a 
foliiferous branch may be seen in a specimen of 8am- 
bucus 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 speci- 
men by the Rev. W. Newbould. 

Jacquin figures an analogous case in Sempervinim. 
sediforme, 1 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, Ta/raxacwiiL, some- 
times offers an illustration of this, and also the daisy 
(Bellis)? 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 ordi- 
nary 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, 

1 ' Misc. Austriac. ad Bot.,' vol. i, p. 133, t. 5. 

- See also Carriere, ' Revue Horticole,' 1866, p. 442 ; and as to pears, 
Radlkofer in ' Bericht iiber die Thatigkert dev Baierischen Gartenbau 
Gesellschaft,' 1862, p. 74, t. i. 



or any absolute increase in fche number of organs, as 
in Prolification. Such ;i case is represented in fig. 
78, which shows a portion of the stem of a species of 
I aleriana, bearing at fche summit, no1 an inflorescence, 
bul a tuft of leaves without the slightest indication of 

Drs. Booker and Thomson relate that in Northern 
[ndia the flowers of Anemone rwularis arc very generally 
absent, and their place supplied by tufts or umbels of 
leaves. 1 In the collection of the late Mr. N. B. Ward 

Fig. 78. — Tuft of leaves replacing the innurescence iu a species of 

was a specimen of lupin in which the flowers were all 
absent, and their place supplied by tufts of leaves. 

A similar appearance has been noticed in Compositce, 
and I owe to the kindness of Professor Oliver the com- 
munication of a specimen of a species of Bidens from 

1 • Flora Indica,' p. 23. 


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 ' Linnasa,' 1837, p. 301, 
Von Cesati figures and describes an analogous case in 
Garduus crispus. The same author 1 records a similar 
instance in the umbel of Seseli coloration, 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 blos- 
soms 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 BryophyUum 
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 inflor- 
escence more than usually branched and covered with 
little tufts of bracts, without a trace of true flowers. 
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 is 
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 
1 ■ Linnsea,' xi, p. '■>* >1. 

\i>\ i:\ I'lTioi g i.i:\\ BS- -Si \i.i:s. 


preserves its usual catkin-like shape, bu1 the flowers 
are replaced by little tufts of Leaves. M. Germain de 
Sainl Pierre mentions a case wherein the flowers of 
Alisma pamassifolia were completely replaced by leaf- 
buds. 1 

Eere, also, may be mentioned the curious aggrega- 
tions of scales which occur in some grasses, in Resti- 
acece, Juncacece, and other orders, in which the inflor- 

Fig. 79.— Spikelets of 
Willeclenovia, composed 
entirely of scales to the 
exclusion of flowers. 

Fig. 80.— Rose Willow, Salix. sp. 

•Bull. Soc. Bot. Fr.,' 1856, p. 53. 


escence 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. 1 

" Rose willows" (fig. 80) owe their peculiar appear- 
ance 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 con- 
tinuance rather than of the arrest of vegetative develop- 
ment. 2 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. 3 

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 palea3 are wholly unchanged, while 
there is no trace of squamulse or of stamens and pistils 
within them, but in their place is a small shoot with 
miniature leaves arranged in the ordinary manner. 

1 ' Trans. Linn. Soc.,' xxvi, p. 142, tab. iv, B. 

2 " Si arbusculam, quae in olla antea posita, quotannis floruit et 
fructus protulit, deinde deponamus in uberiori terra ealidi caldarii, 
proferet ilia per plures annos multos ac frondosos ramos, sine nllo 
fructu. Id quod argumento est, folia inde crescere, unde prius enati 
sunt flores ; queinadmodum vicissiin, quod in folia nunc succrescit, id, 
natura ita moderante, in flores mutatur, si cadein arbor iterum in olla 
seritur." — Linnaons, ' Prolepsis,' § iii. 

3 < Rev. Hortic,' May, 1808, * Gardeners' Chronicle,' 1868, pp. 572, 737. 

\ i\ [PAR01 s PLANTS. 


The grasses most commonly affected in lliis manner 
are Dactylis glomeratal, I'o'i bulbosal, Poa a/rvrma /, 
/'. fyriviali&I, prat&nsis /, alpi/na!, angvstifolia, and /a"', 

Fig. 81. — Portion of panicle of Alva vivipara and separate floret. 

Cynosurus cristatus, Festuca ncmoralis, F. ovina!, Ghj- 
ceria fluitans ! ', Gl. aquatica, Aira alpha!, ccespitosa!, 
PI ileum phalaroides, Lolium perenne !, Alopecurus pra- 
tensis !, Agrostis alba, Holcus mollis ! 

From an examination of the structure of viviparous 
grasses Yon Molil 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 sur- 
rounds the stem or axis of the spikelet entirely, and 
both its margins cohere towards its lower extremity. 1 

A similar condition occurs not infrequently in 
Polygon/urn vwipa/rum, and in Jimcacece, Cyp&racece, &c. 

1 Cited in ' Annals Nat. Hist.,' 1845, vol. xv, p. 177. 


In the genus Alliwm an analogous formation of little 
buds or bulbils takes place in lieu of flowers ; this is 
specially the case with A. vmeale, 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 

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, 1 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 Nym/phcea guinea nsis, some Arads, &c. 
The same condition has been met with as a teratological 
occurrence in the leaves of Cardamine pratensis, Hya- 
riutliiis I'nii;:<Jsii ', Drosera intermedia? AraMs pwmila, 
Ghelidonw/m majus, Chirita sinensis, 3 Ejnscia tricolor* 
Zarn ia, &c. 5 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. phyllo- 
maniaca, Mart., is the species best known as manifest- 
ing this tendency, but others have it also/' 

Buds are also very often formed upon the margins 
of the leaf, the best known instance of which occurs in 

1 'Ann. Scienc. Nat.,' vol. xiv, 1860, p. 13. 

- Naudin, ' Ann. Sc. Nat./ 2nd ser., 1840, vol. xiv, p. 14, fig. 6, pi. i 
(Drosera). St. Hilaire, ' Comptes Rendus,' ix, p. 437. 

3 Hance, ' Hook. Joum. Botany,' 1849, vol. i, p. 141, pi. v. 
* Booth, ' Gard. Ckron.,' Jan. 1st, 1853, p. 4. 

5 Lindley, ' Theory of Horticulture,' ed. 2, p. 273. 

6 ' Hook. Joum. of Botany,' 1852, iv, p. 206. See also the curious 
Begonia gem/mipara, 'Hook. fil. Illust. Himal. Plant..' t. xiv. 

Al>\ i:\IITlMl v |;| | )S> 


ophylhvm eah/cmum ; Weinmann 1 figures an instance 
of this kind in Alchemilla minima, or they may occur 
upon the lower surface of the leaf, as in OrnithogaVwm 
scilloides and longe-bracteatum. M. Duchartre 2 men- 
tions 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 Drost ra before mentioned. 

Fig. 82. — Formation of shoot on leaf of Episcia bicolar. 

Gardeners occasionally avail themselves of this for- 
mation 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 

1 • Phytantk.,' n. 36, d. 
"Ann. Scienc. Nat.,' 3rd serii -. l v ">:;. vol, xix. p. 251, tab. I I 



normal condition in the curious Btangeria paradoxal 
and Mr. Berkeley records an instance of this in sea- 

Fig. 83. — Adventitious buds in sea kale. 

kale 2 (fig. 83) where the crown had been injured, and 
buds were seen sprouting from its centre. 

It will be remarked that the adventitious production 

Fig. 84— Hyacinth bulb cut Fig. 85.— Showing the formation 

across to induce the forcnation of of new bulbs on the cut edges of 
new bulbs. an old hyacinth bulb. 

Carriere, ' Revue Horticole,' 1868, p. 184. 
• Gard. Chron.,' 1858, p. 556. 

\e\ BNTITI0U8 Bl DS. 


• 'I' buds, like i lint of roots, is very often consequent on 
decay or injury. The Dutch bulb-growers have availed 
themselves of tliis latter circumstance in the propa- 
gation of hyacinths. Mr. Fortune, who published 
some articles on this subject in the 'Gardener's 
Chronicle,' 1 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. 8 1 , 
85). The other method is effected by scooping out 
the interior of the base of the bulb, thus leaving ex- 
posed the cut ends of the sheathing leaves arranged 
concentrically; along these lines the new bulbs are, 
after some time, formed in great numbers (fig. 80). 

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 

Production of gemmae in place of spores. — An instance of 
18G3, p. 556, &c. 



this is recorded by Dr. Montagne 1 in the case of a 
moss, Encamptodon perichcetialis, in which, in the in- 
terior of the capsnle, in lieu of spores numerous minute 
gemma3 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 Busciis, in Helwingia, Erythrochiton 
hypophyllantlms, 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 con- 
venience sake it is placed in this section, as it is un- 
certain at present where it properly belongs. 

This curious plant has been described and figured 

Fig. 87. — Three- lobed end of 
outer palea of Nepaul barley 
bearing supplementary florets. 

Fig. 88. — Three spikelets of 
Nepaul barley. 

■ Ann. Nat. Hist.,' 1845, vol. xvi. p. 355. 

\i>\ ENT1TI01 s n.uw ERS. 

i ■> 

l)\ hniiscli in the l-ltli volume of the ' Linnaea,' p. L24, 
fc.iv; also by Professor Ilenslow, 'Hooker's Journal 

of Botany, 5 L849, vol. i, p. 33, tabs. 2, 3. The lower 

palea of this plant forms an inverted flower-hud upon 
its midrib. In sonic fresh specimens which J have 

Fig. 89. — Lip of outer palea 
of Nepaul barley. 

Fig. 90. — Supplementary rachil- 
lus or outer palea of Nepaul bar- 
ley bearing florets. 

Fig. 91. — Diagram sbowing ar- 
rangement of supplementary ra- 
cbillus and florets. 

Fig. 92.— Supplementary floret 
of Nepaul barley ; palea removed. 

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 paleaa 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 palea 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 ex- 
ceptional a case as that of the Nepaul barley. 1 

Production of flower -buds in place of leaf -buds. — Under natu- 
ral 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, 2 
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 

1 See also Lindley, ' Veg. Kingd.,' p. 109 et 116a, where the views of 
Raspail, R. Brown, Mohl, Henslow, and others, are discussed. 

8 It has been observed that if a plant is supplied with copious nourish- 
ment the flowering-period is delayed ; but that moderate or even scanty 
nourishment accelerates it. Goethe, ' Metam.,' § 30. See also Wolff, 
'Theoria Generationis,' 1759; Linn. ' Prolepsis,' §§3 and 10. 

HETEE0TAX1 . I i < 

supply of Qutriment, and other means all based on the 
same principle. Some of the Cape bulbs (Cyrtanthus) 
are known not to produce fcheir flowers till their leaves 
have received, in some manner, a check. Fires which 
often destroy the herbage thus have the effect of throw- 
ing 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 con- 
tracted 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, 18-58, 
p. 316) a branched spine of Gleditschia bearing a flower 
at the end of each of the subdivisions. 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 
1 Gardeners' Chronicle' for 1865, p. 769, by the Rev. 
M. J. Berkeley, who describes the formation of a 
flower-bud on the surface of a petal of Glarkia 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. Avery 
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. 1 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. 2 Further 
reference to these fruits will be made under the head 
of Hypertrophy. 

M. Trccul 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. 3 A similar growth 
was observed in the fruit of PMladelphus sjn'riosushj 
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. 4 

The fruits of Opuntia Salmiana, 0. fragilis, 5 0. mona- 
cantha, and of some species of Echmocactus, 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, 

1 Moquin-Tandon, p. 384; also Lindl., 'Elements of Botany,' p. 65, 
fig. 130 ; " Theory of Horticulture," p. 86. ' Gard. Chron.,' 1851, p. 723 ; 
[rmish, 'Mora,' L858, p. 38, &c. 

2 Caspary, ' Bull. Soc. But. Fr.,' vol. vi, 1859, p. 235; also Payer, ibid., 
vol. i. 1854, p. 283. 

3 Trecul, ' Aim. Se. Nat.,' 2nd ser., \ol. xx, p. 339. 

4 'Bull. Soc. Bot. Fr.,' vol. rii, L858, p. 331. 

5 'Bull. Soc. Bot. IV.,' vol. i, p. 306, vol. v, p. 115. ' IUustr. Hortic.,' 
srii, L865, Misc. 7;'. -Rev. Horticole,' !*<'>o. p. 204, et 1867, p. UJ. 



and then a circle of small )>u<ls, like those of the stem, 
may be seen at the top of the fruit, each hud springing 
from the axil of a little tuft of wool and -pines found 
• »ii the fruit. TheBe little buds clone-ate 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. Thespecimen from which the figure (fig. 
!»:')) 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 

Fig. 93. — Small buds projecting from the edges of the fruit in OpwnMa. 

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 Pereshia, which bear leaves on their sur- 
face arranged spirally ; indeed, the fruits of Peredia 
Biro are mentioned as producing buds from their 


summits, in the same way as the Opuntia just cited. 
P. Bleo is said, by M. Delavaud, 1 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 
cither 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. 2 

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 Yon 
Mar this, 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 Pomacece, and Baillon 3 succeeded in producing 
buds on the surface of the inferior ovary of Jussicea. 

Some of the cases just mentioned have been con- 
sidered to be instances of prolification of the fruit, 
but the fruit has little to do with the appearances in 

Formation of adventitious flowers and fruits within the ovary. — 
This generally arises either from substitution of a 

' ' Bull. Soe. Bot. Ft.,' 1858, p. 685. 

2 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, Ft.,' vol. v, 1855, 
p. 072. Seringe et Heyland, ' Bull. Bot.,' i, p. 8. ' Pallas Enuni. Plant. 
Hort. Demidoff,' append, c, ic. 

3 ' Adansonia,' i. 181. 


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 prolifi- 
c-it ion strictly. 

The cut, fig. 94, represents a case where, in the 
dilated upper portion of the ovary of Sinapis a/rvensis, 

Fig. 94. — Distended pod of Sinapis arvensis bearing in the interior 
stalked iiower buds. 

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 aw/phibiwm, Brassica 
Bapa, and Passiflova quadrangularis. 

In Bromfielcl's ' Flora Vectensis,' p. 35, the following 
account is given of an abnormal development in Carda- 
mine 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 hi 
the pods above had burst from their confinement and 
appeared as semi-double fiow r ers ; the valves of the pod 
answering to the true calyx. * * * From their verti- 
cillate arrangement it is evident that these petaloid 
expansions w^ere not transformed seeds, but simply a 
development of the common axis wdthin 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 1 also records a 
case of the same nature in Sina/pis arvensis. 

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 

Fig. 94. — Portion of the interior 
of the silicle in Cheiranthtis Cheiri, 
showing adventitious pod in the 
place of an ovule. 

Fig. 95. — Adventitious pod from 
fig. 94, enlarged. 

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 Bceckea diosm-ce- 

1 ' Adansonia,' vol. iii, p. 351, tab. xii. 



folia, Rudge, for an examination of which he is in- 
debted to Mr. Ben1 ham. 

Fig. 96. — Section of Barbarossa Fig. 97. — Grape with supple- 

grape showing adventitious grape mentary fruit in the interior. 
in the position of a seed. 

Id 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 adhe- 
rent, is surmounted bya 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 con- 
tinuous with that organ, show externally any indica- 
tion 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 fila- 
ments, 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 carpellarv 
leaves) . 

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 Bceckea there were perfect stamens proceeding 

Fig. 98. — 1. Vertical section of flower of Bceckea diosmcefolia, showing 
stamens within the ovary ; magnified ten times. 2. Transverse section of 
ovary. 3. Stamen. 4. Imperfect stamen. 


1 85 

from perfect and completely closed ovaries. Moquin- 
Tandon 1 cites from A.gardh an instance which seen, 
more closely to resemble the state of things in the 
Bceckea, and which occurred in a double hyacinth, 
wherein both anthers and ovules were borne on the 
same placenta. Probably 3 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 BcecJcea nothing at all unusual was ob- 
served 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 Babmg- 
tonia s a prolongation of the placenta.' 

Formation of pollen within the ovules. — This has now been 
recorded in two instances by Mr. S. J. A. Salter in 
Passiflora ccerulea and in P. pahnata* and by the author 
in Rosa arvensis. 41 

In the case of the passion-flower there were various 
malformations in the ovaries, which were all more or 

Fig. 99. — Pollen within the ovule of Passiflora (after Salter). 

1 « Elein. Terat. Veget.,' p. 218. 

J Masters, ' Journ. Linn. Soc.,' vol. ix, 1866, p. 334. 

3 ' Trans. Linn. Soc.,' vol. xxiv, p. 143, tab. xxiv. 

4 'Brit. Assoc. Report,' Dundee, 1867; and Seemann's 'Journal of 
Botany,' 1867, p. 319, tab. lxxii. figs. B 1—9. 


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 ordi- 
nary 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, con- 
taining, 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 

P0LLINIFER01 S OV1 l.l'.s. L87 

of the ovule, while the pollen was formed within the 

nucleus. In no case was any trace of embryo sac 
to be s» 'en. 

The main interest, as Mr. Salter remarks, in these 
cases is physiological; so far as structure alone is con- 
cerned, 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 indi- 
viduality 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 abso- 
lute form of hermaphroditism." 

It must, however, be remarked that the term sub- 
stitution 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 con- 
sidered 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 per- 


ceptible. 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 cir- 
cumstances 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. 

Honiomorphic flowers of " Conrpositse." — In a large section 
of the Composite^ 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 regu- 
larity 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 Liguliflorcr, 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 in- 
florescence, such as the change from a definite centri- 
fugal form of inflorescence fco 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 pro- 
perly 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 tw r ice in the 
same season; this usually arises from the premature 
development of buds which, under ordinary circum- 
stances, 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 Equisetacece, 1 in Restiacece, and other orders. In the 
equisetums, the condition in question has been specially 
noticed to occur after prolonged drought. 

Equisetacece 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 cat- 
kins of Salix cinerea were placed at the ends of the 

1 Duval Jouve, ' Hist. Equiset. France,' L8ti4. p. 154. 


branches instead of being lateral productions ; more- 
over the usual articulation was not formed, so that 
the catkin was persistent instead of deciduous. 1 

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 some- 
times 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 oppo- 
site to the maro'inal crenatures. But it is also fre- 
quently 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-sori- 
ferous condition. Among the ferns in which this 
condition has been observed are the following; : Seolo- 
pendrium vulgare, Polypod/iwn amymalv/m, Hook., As- 
plenium Trichomanes, Gionidium Moor&i} 



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 habi- 

1 ' Flora,' t. xxiv, 1841, p. 340. 

2 Moore. - Xatnre-Printed British Ferns. ' 8vo edition, vol. ii. p. 135, 
tab. lxxxv, b. Ac. 


tually suppressed, a tendency towards structural com- 
pleteness rather than the reverse. It musl be also 
understood thai the following remarks apply to struc- 
tural points only, and are not intended to include 
the question of function. The occurrence of hetero- 
morphic unions renders it necessary to keep in mind 
thai 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 thai 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 inflor- 
escence 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. 1 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 Coniferce 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 

1 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. 



in July, 1860, some malformed cones of Ahies excelsa, 
in which the inferior part of the axis was covered with 
stamens, whilst the terminal por- 
tion 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 some- 
what modified, the anther cells 
being diminished, whilst the 
scale-like crest had become more 
elongated and pointed, in fact, 
more or less resembling the or- 
dinary bracts. 1 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. 

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. t>5. Caspary, ' De Abietin. flor. fern. 

struct morphol.' Schleiden, ' Principles,' English edition, p. 299. Mohl, 

'Verm. Schrift,' p. 45. Meycn in ' Vfiegin. Archiv.,' 1838, p. 15$. 

Cramer, ' Bildungeabweich, 5 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. 

1 Professor Dickson concludes from the examination of these struc- 
tures 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, 



Change from the monoecious to the dioecious condition. — This 
is of less frequent occurence than might hare been 
anticipated. In the 'Gardeners' Chronicle,' 1817, 
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.— Andro- 
gynism. — This is of far more common occurrence than 
the preceding. 

Fig. 101. — Monoecious inflorescence of Hop. 

In the hop (Humuhs 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 bvhini with monoecious inflorescence: 
Schismatopera distichophylla, Mozinna peltata, Hermesia 
castaneifolia. Oliver mentions (' Hook. Icon. Plant.,' 
t. 1044) that in Leitneria fioridana 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 Oarex, while in the case of Salix r&pens, 
Hampe 1 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 2 says that a plant of Stavntonia 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 nut- 
meg is also said to vary in sexual characteristics from 
time to time. 3 In addition to the genera, already 
named, in which this production of flowers of both 
sexes has been observed may be mentioned Tax us ! 
Gurmera ! Urtica! Mercit/rialis ! Bestio ! Cannabis! 
Salix ! Humvlus ! as well as others in which the 
change is less frequent. 

Among cryptogams a similar change occurs. As an 
illustration may be cited Levcobryvm gigantewm, as 
cpioted from Midler 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 

1 ' Linnaea,' xiv, 367. 

2 ' Rev. Hortic.,' January, 1867. 

3 See Royle, ' Man. Materia Medica,' ed. 1, p. 567. 

i \is i:\r.\UTV. [95 

Berol.,' ii, p. 259. Tausch, ' Bot. ZeiV 1833, i. p. 229. Koch, ' Synops. 
Hor. Germ.,' 740. Host, 'Hor. An at.,' Li, p. till [B.mirabU 
also Eegelmaier, ' Wtirttemberg Natorwissenschafl Jahreahefte,' I-';*'.. 
]>. :!<>. Other referencea t" leaa accessible works are given in ' Ldnnaea,' 

xiv, i>. 372. 

Change from hermaphroditism to unisexuality. — Many 
flowers ordinarily hermaphrodite as to structure, be- 
come unisexual by the abortion or suppression of 

their stamens, or of their carpels, as the case may be. 
This phenomenon is lessened in interest sin Ce the 
demonstration of the fact by Darwin and others, thai 
many plants, structurally hermaphrodite, require for the 
full and perfect performance of their functions the co- 
operation of the stamens and pistils, belonging to dif- 
ferent individuals of the same species. 

Some of the Rcmunculacece constantly exhibit a ten- 
dency towards the dioecious condition, and the rarity 
with which perfect seeds of Ranunculus Mcaria 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. Speci- 
mens of Ra/wwrbculus 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 pro- 
duction of stamens, wdiile 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 acci- 
dentally prevented from perfecting their female organs. 1 

Mr. Darwin 2 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 de- 
veloped in more southern districts. 

Honclcenyapeploides affords another illustration of the 
sexual arrangements in the flower being altered as it 
would seem by elimatal 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. 3 

Treviranus 4 says that the flowers of Hijijinris 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. 

1 Thomson, ' Gardener's Assistant,' p. 577. 

2 ' Variation of Animals and Plants,' i, 353. 

3 Babington. ' Ann. Nat. Hist.,' vol. ix. 1852. p. 156. 

4 • Plvys. dor Gewachse,' ii. p. 323. 


Change from unisexuality to hermaphroditism. This Occur- 
rence depends on one of two causes, either organs are 
developed (stamens or pistils as the case may b 
winch arc habitually absenl in the particular flower ; 
or some of the stamens may be more or less com- 
pletely converted into or replaced by pistils, or vici 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, inas- 
much as certain organs which habitually undergo 
suppression at a certain stage in their development, 
by except ion, 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 leasl 
structurally. I have myself seen hermaphrodite 
flowers in Oucurbita, 1 Mermrialis, Cannabis, Zea Mays, 
and Aucuba japonica, as well as in many Restiacece, 
notably Gannamois virgata and Lepyroclia Jierinajjhro- 
dita. Spinacia oleraeea, Rhodiola rosea, Cachrys taurica, 
and Em/petrwm nigrum are also occasionally herma- 

Gubler 2 alludes to a similar occurrence in Pistada 
LenUscus, wherein, however, he adds that there was a 
deficiency of pollen in the flowers. 

Schnizlein 3 observed hermaphrodite flowers in the 
beech, Fagus sylvatica, the ovaries being smaller than 
usual, and the stamens epigynous. 

Baillon 4 enumerates the following Ev.jjhorbiacect as 
having exceptionally produced hermaphrodite flowers, 

1 See also Schlechtendal, ' Linmea.' viii, p. 623, and Lindley. ' Veg. 
Kingd..' p. 315. 
1 • Bull. Soe. But. France,' vol. ix. p. 81. 

3 Cited in Henfrey. ' Bot. Gazette,' 3, p. 11. 

4 Baillon. 'Etudes du Groupe des Euphorbiacees,' p. 2<V>. tab. xv. 
fig. 19, tab xix. tig. 31. 

J 98 


Crozoplwra tinctoria, Suregada sp., PhyllantJms longi- 
folius, Breynia sp., Philyra brasiliensis, Bicinus com- 
munis, Conceveiba macrophylla, Gluytia semperjlorens, 
Wall, non Boxb. Mercurialis annua and Cleistanthus 

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. 1 

Robert Brown 2 observed stamens within the utricle 
of Car ex acuta, and Gay is stated by Moquin (' El. Ter. 

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. 

1 See also Guillemin, ' Mem. Soc. Nat. Hist. Paris,' I, p. 16 ; herma- 
phrodite flowers in Euphorbia esula, 

2 ' Prod. Flor. N. Holl.,' p. 242. 



Veg., J p. 8 1-3) to have observed a similar occurrence 
in ( ',!,■> b glauca. 

Paascli 1 observed a similar occurrence in G. ccespitosa, 
and Schauer, in C. pcdudosa* though in the latter in- 
stance the case seems to have been one of transfor- 
mation or substitution rather than one of hermaphro- 

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. 3 

In Begonia frigida* 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 

Fig. 103.— Hermaphrodite flower of Carioa Papaya. 

from the former, but simply to the presence of ovaries 
where, under ordinary circumstances, stamens only are 

1 ' But. Zeit..' 1837, p. 335. 

1 ' Prianz. Terat.,' von Moquin-Tandon, p. 208. 

3 Schnizleiu, loo. cit. 

4 'Bot. Mag.,' tab. 5160. fig. 1. See also ' Gard. Chron.,' 1860, pp 
116,170; 1861, p. 1092. 



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 

In hermaphrodite flowers of Garica 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 

Fig. 104. — Ovuliferous anthers — Cucurblta. 

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 : 





















That of the hermaphrodite blossoms would be, in brief, 
as follows : 


One of the most curious cases of this kind recorded 
is one mentioned by Mr. Berkeley, 1 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, pollini- 
ferous, 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. 



The deviations from the ordinary direction of organs 
partake for the most part more of the nature of varia- 
tions 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 

1 ' Gard. Ohron.,' 1851, p. 499. 


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 result- 
ing from the natural growth of the plant. These and 
other causes tend to alter the direction of parts very 

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 Tray a natans, the roots of which 
in germination are directed upwards towards the sur- 
face of the water. So in Sechium edulc, 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 some- 
times even by seed ; hence the origin of fastigiate 
varieties of elms, oaks, thorns, chesuuts, and other 
plants which may be met with in the nurseries. 


Sometimes when the top of the main stem is de- 
stroyed by disease or accident, one of the heretofore 
lateral shoots takes its place, and continues the de- 
velopment of the tree in the original direction. It is 
often an object with the gardener to restore the sym- 
metry of an injured tree so that its beauty may ulti- 
mately not be impaired. 1 

Climate appears sometimes to have some influence 

1 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. 
Northnanniana, 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 detri- 
mental 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 illus- 
trative 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. Loicii 
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 


on the direction of branches, thns Dr. Falconer, as 
quoted by Darwin, 1 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. 2 

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 

1 'Variation of Animals and Plants,' ii, p. 'J77. 
- Quoted in ' Gard. Chron.,' 18(57, p. t>54. 



then elongate into a shoot and no cavity is formed. A 
fig observed by Zuccarini (figs. 105, JOG) appears to 
have been formed in a similar manner, the flower-bear- 
ing 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. 

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 maybe 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 

Fig. 105. — Fig showing pro- 
longed inflorescence and projecting 

Fig. 106. — Section of the same. 

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 tlieMuseum 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. 

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 

Fig. 107. — Hollow turnip, showing some of the leaves inverted and 
occupying the cavity. 

some of these movements bear to the process of ferti- 
lisation, 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 accom- 



pany 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 indi- 
vidual members, for instance of Gloxima, 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. 

Fig. 108. — Flower of normal Gloxinia. 

Fig. 109. — Flower of Gloxinia, erect and regular (regular Pelorin). 

Fig. 108 shows the usual irregular form of Glosdnia, 
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. Ill are 
shown the stamens and style of the two plants respect- 
ively. 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 rela- 
tive 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 con- 
genital 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. 

Fig. 110. — Calyx of erect 

Fig. 111. — Stamens of erect 
regular, and of pendent irre- 
gular-flowered Gloxinia. 

A similar alteration accompanies this form of peloria 
in other flowers (see Peloria). A change in direc- 
tion 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 benl downwards towards the stunted 
carpels instead of being, as they usually are, straight. 
Amongsl orchids, where the pedicel of the flower or 
the ovary is normally twisted, so thai the labellum occu- 
pies the anterior or inferior part of the flower, it fre- 
quently 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 

Fig. 112. — Flower of Azalea, showing the corolla reflected. 

young flowers of this singular variety have not been ex- 
amined 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. 

Moquin 1 mentions a flower of Rosa alp'uta 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 libe- 

1 Loc. eit., p. 315. 




rating the style and allowing it to assume a vertical 

M. Carriere 1 lias 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 

Fig. 113. — Flower of Cuphea miniata enlarged, showing protrusion 
and hypertrophy of an erect placenta, after Morren. 

Fig. 114. — Placenta from the flower shown at fig. 113; the ovary is 
membranous and torn, the placenta, erect and ovuliferous, after Morren. 

tree as usual, while the smaller one was pointed in 
exactly the opposite direction. The larger fruit had the 

i ' Rev. Hortic.,' 1868, p. 110. 

.11 \\i,|. 01 DIRECTION. L*l 1 

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 cir- 
cumstances ; 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 Gissus? 

Morren describes a curious condition in some flowers 
of Cv/phea miniata, in which the placenta protruded 
through an orifice in the ovary, and losing the hori- 
zontal direction became erect (figs. 113, 114). A similar 
occurrence happened in Lobelia ervn/us. To this con- 
dition the Belgian savant gave the name of gymnaxony. 2 

1 Plancbon and Mares, ' Ann. Sc. Nat.,' 5 ser., torn, vi, 1866, p. 228, 
tab. xii. 

' ' Bull. Acad. Belg.,' xviii, part ii, p. 293. 



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. 1 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 sup- 
posed that all flowers, no matter how irregular thev 

1 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 symmeti'ical 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 aiTangement, form, and number of its 
component elements. 


subsequently became, began by being strictly symme- 
trical or regular, and that subsequent alterations were 
produced by inequality of growth or development. 
The researches of organogenists have, however, dis- 
pelled 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. 1 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 know- 
ledge, to determine whether a given arrangement is 
congenital or acquired subsequently to the first de- 
velopment, whether for instance an isolation of parts 
be due to primordial separation or to a subsequent dis- 
union of originally combined organs, see p. 58. With 
reference to the changes in the form of organs, how- 
ever, it is in general more easy to ascertain the proxi- 
mate 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, irre- 
gular development; hence the use of the following 

1 See Baillon, ' Adansonia,' v, 17<i. 



terms — Stasimorphy, Pleiomorphy, Metamorphy, and 
Beteromorphy — to include teratological changes really 

or apparently due to one or other of the causes above 
mentioned. The classification here adopted is ofcour.-r 
to a considerable extent an arbitrary one and subjeci 
to correction or modification, as the knowledge of the 
developmenl of the flowers in tin' various genera of 
plants advances. 



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 "develop- 
ment " is checked. Such conditions may even be pro- 
pagated 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 ante- 
cedent condition. Stasimorphic changes affecting 
principally the relative size of organs — such, for in- 
stance, 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. 

1 traffic- fiorxpuHjiQ. 





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 satis- 
factorily, while the reader may be referred for other 
instances of like nature to the sections on Peloria, 
Atrophy, Suppression, Dimorphy, Substitutions, &c. 

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 

Fig. 115. — Juniperus sinensis. Two forma of leaves on bran 
the same shrub. 


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. 1 Most of these cases are elsewhere referred 
to in this volume under the particular form of mal- 
formation 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. 

1 See a paper of Professor C. Morren's on " Floral Stesoniy " in 
' Bull. Acad. Belg.,' t. xix, part ii, p. 519. 





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 forma- 
tion of irregular parts in increased number, so that the 
symmetry of the flower is rendered perfect, as in the 
original peloria of Linnseus, 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. 1 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 

Fig. 116. — Regular Peloria, Fig. 117. — Sepal, petal, &c, of 

Delphinium. regular- flowered Delphinium. 

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 

1 " On the existence of two forms of Peloria," by M. T. Masters. 
' Nat. Hist. Review,' April, 1863. 



in other species of this genus. Baillon, 1 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 in- 
creased 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 occur- 
ring 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 

Fig. US. — Regular peloria, Fig. 119.— Double Violet, flower 

Viola. regular, petals multiplied, stamens 

and pistils petaloid. 

1 Baillon. ' Adansonia.' iv. p. 149. 



hairs on the lateral petals; 4, equal stamens whose 
anthers are sometimes entirely destitute of the pro- 
longed 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. 1 Between 
these cases and the ordinary spurred forms as well as 
those with an increased number of spurs, many inter- 
mediate 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 sub-division of the order 
(Alsodeice) 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 circum- 
stances, 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 times partially. The direction of the stamens 
and style, and even that of the whole flower, becomes 

1 Similar cases are figured in 'Hort. Eystettens. Ic. PL Vera.,' fol. 4, 
f. 1, 2. Viola martin multiplici flore. 



altered from the inclined to the vertical position. In 
addition to these changes, which are those most com- 
monly 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 Trojxeolum 
majus in which the flowers were perfectly regular and 
devoid of spurs, 1 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 Papilionacece the Laburnum 
and others have been noticed to produce occasionally a 
perfectly regular flower in the centre, or at the ex- 
tremity 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 Gesneracece, Bignoniacece, Scroj)hu- 
lariacece, and other families of like structure, regular 
peloria is not uncommon. Fig. 120 represents a case of 

Fig. 120. — Regular peloria, Eecrenwcarpus scaber. 

this kind in Eccremocarpus scaber, conjoined, as is fre- 
quently the case,' with dialysis or separation of the 
petals. 3 Many of the cultivated Gloxinias also show 

i ' Linna3a,' 1837, p. 128. 

2 M. Bureau, 'Bull. Soc. Bot. Fr.," ix, p. 91, describes two genera of 
Bignoniacece in which the flowers are normaUy regular and six parted. 



erect, regular, five stamened flowers, but these are 
probably cases of irregular peloria. 

A solitary flower of Pediculcuris 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. 1 

The passage of ligulate to tubular corollas among 
Composites is not of such common occurrence as is the 
converse change. I owe to Mr. Berkeley the communi- 
cation of a capitulum of a species of Bidens, in which 
there was a transition from the form of ligulate co- 
rollas to those that were deeply divided into three, 
four, or five oblong lobes. These then were instances 
of regular peloria. 

Fig. 121. 


-Flower of Cattleya marginata. Lip replaced by a flat 

In Orchidacece a similar change is not by any means in- 
frequent ; in a few, indeed, a regular flower is the normal 

1 See ' Trans. Linn. Soc.,' vol. x. p. -227 


character, as in Dendrobium normale, Oncidium heteran- 
tJtum, Thelymitra, etc. Fig. 121, reduced from a cut in 
the 'Gardeners' Chronicle,' 1854, p. 804, represents an 
instance of this kind in Gattleya marginata. 

From the same journal the following account of 
a case of peloria in Phalcenopsis 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. Pro- 
fessor Reichenbach also exhibited at the Amsterdam 
Botanical Congress, of 1865, a flower of Selendpediuub 
caudatum with a flat lip. 

M. Gris 1 has placed on record some interesting cases 
of peloria of this kind in Zingiber zerumbet ; in the more 
complete forms the andrcecium 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 

Under this head may likewise be mentioned those 
cases in which the normal, or at least the typical sym- 
metry of the flower is restored by the formation of 
parts usually suppressed ; thus Moquin cites an abnor- 
mal flower of Atriplex* 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 Boccouin with a 

1 'Ann. Sc. Nat.,' ser. 4, 1859, torn, xi, p. 204, tab. 3. 

2 'El. Ter. Veg.,' p. 342. 


corolla. Arn m maculatum has likewise been met with 
provided with a genuine perianth as in Acorus and 
other Orontiads. The unusnal development of the 
Bexnal organs in diclinous flowers has been alluded to 
under the head of heterogamy, and other cases wh< 
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 petal- (Antirrhinum, Verbascum, 
I , 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 Tropceolum, Viola, &q. 
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 
of the flower. 

From overlooking the occasional existence of this 
form of peloria, new genera have sometimes been 
formed on insufficient grounds. The genus Aceran- 




thus, for instance, consists of species of Epimedium in 
which the customary spurs are not formed. 1 

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. 2 

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 ! 

Epimedium, sp. 
*Pelargonium zonale ! 

inquinans ! 

Tropseoluni majus ! 
*"Wistaria sinensis. 

*Cytisiis Laburnum ! 

Trif olium repens ! 
*Conipositse, gen. pi. ! 

Lonicera, Periclynienum ! 

Streptoearpus Rexii. 
*Digitalis purpurea. 
*Scrophularia aquatica. 
*Linaria vulgaris ! 
-"Antirrhinum majus ! 

Verbascuru nigrum ! 

Columnea Schiedeana. 

Halenia heterantlia. 

Galeodolon luteum. 

Prunella vulgaris ! 

Salvia, sp. ! 

Teucrium campanulatum. 

Betonica alopecuros. 

Eccremocarpus scaber. 

1 Marcband. ' Adansonia,' vol. iv, p. 127. 

- ' Bull. Acad. Belg..' xvii. p. 17. " Fuchsia." p. 169. 


Pedicularia Bylvatioa A triplex, ro. 

Zingiber Zerumbet. CattLya Mnssi;i ! 
PhausnopaiB amabilis! mar&inata. 

Phakenopsis Schilleriana. Calantlie vest it a ! 

Haliciiaria. Oneidium, sp. ! 

*Orchis niorio. Selenipediuni caudatum. 

masciila. Aruui niaculatum. 
*Dendrobiuiu, sp. . 

In addition to the references already given, further 
information on this subject may be gained from con- 
sulting the following publications. See also Irregular 

Giraud, ' Bot. Soc. Edinb.,' Dec. 12, 1839, Antirrhinum. Dareste, 
' Ann. Sc. Nat.,' ser. 2, 1812, 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 aber- 
rantes et regukvrisantes.' Reichenbach fil. ' De pollinis orchid, genesi 
ac structural 1852, Oneidium. Clos, 'Mem. Acad. Toulouse,' vi, 1862, 
Salvia. Caspary, ' Yerhandl. Phys. (Ekon. Gesell. Konigsberg,' I860, 
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 regidarite 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. 



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. 



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 
TJpsal. This was considered so marvellous a circum- 
stance that the term peloria, from the Greek 7rAwp, a 

1 nA{iot,'-/*6p$H<j<7i(.. 


prodigy, was applied to it. 1 After a time other irregular 
flowers were found in like condition, and bo 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 Avhere 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 

Peloria is either complete or incomplete ; it is com- 
plete 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 

1 ' A mom. Acad.,' i, p. 55, t. iii (1744) : — Tlie following note refers to 
Linne's notion that these forms were due to hybridization. It is 
extracted from Gmelin's edition of the ' Systerna Naturae,' 1791, p. 931 . 
" Linarice proles hybrida, ejusdemque qualitatis et constans, radicibus 
infinite sese multiplicans charactere fructificationis diversissima, corolla 
regulari, quinque-corniculata, pentandra, ut genus proprium absolute 
constituent et distinctissimum, nisi fructus frequentissime abortiret. 
Naturae prodigium. Ita quideni a Linne. Verisimilior autem videtur 
ea opinio, quae peloriam pro peculiari degeneration e monstrosa floris 
habet, in quam inclinare boc genus (Linaria) prae aliis, similis a forma, 
deflexio in aliis speciebus, e. g. spuria Elatine, cymbalaria, observata, 
. . . . Merk., ' Goett. gel. Anz.,' 1774, n. 121. Linck, 'Annal. 
Naturg.,' i. p. 32." 


serve to show in what way the irregularity is brought 
about. In Antirrhinum, Lmaria, &c, intermediate 
forms show very clearly that it is to the repetition of 

Fig. 122. — Peloric flower of Calceolaria. 

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 ! 

Morren l 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 termi- 
nating in two hollow teeth, between which is the 

1 ' Bull. Acad. Belg.,' xviii, part i, p. 591. Lobelia, p. 137. 


narrow orifice of* the corolla. The colour at the b 
of the tube inside is as in the perfect peloria ; while 
round the summit of the tube, in both cases, the inten- 
sity 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 Gytisus LaJmrTi/u/m, Glitoria 
Ternatea, Trifolium r<'jx-ns, and other Papilionacea > , 
it is the "standard," the form of which is repeated. 
In the case of peloric aconites 1 the lateral and some- 
times the inferior coloured sepals assume the hooded 
form usually peculiar to the upper sepal only, the 
number of the petals or nectaries being correspond- 
ingly increased. Balsams become peloric by the aug- 
mentation in the number of spurs. 2 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, 3 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 direc- 
tion and relative position with reference to the other 
parts of the flower is different from ordinary. This 
was the case in some flowers of Phalcenopsis equestris 
sent to me by Mr. Wentworth Buller. Fig. 123 repre- 
sents a flower of Aristolochia can data with two lips, 
for which I am indebted to Mr. W. H. Baxter. 

From these cases it is evident that the flowers in 

1 See also Seringe, ' Esquisse d'une Monogr. clu genre Aconitnm,' p. 124. 

2 Schlotterbec, ' Act. Helvet.,' t. ii, pi. i, Roeper. Balsam, p. 10, note. 

3 Masters. " Peloria. &c, Ophrys arcmifera" ' Journ. Linn. Soc,,' viii. p. 



question become regular by the repetition of the irregu- 
lar parts. 

It is probable that peloria may occur in any habitu- 
ally irregular flower, and that, if more attention were 
directed to the subject, illustrations might be obtained 

Fig. 123. — Two-lipped flower of Aristolochia cauclata. 

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 loth, 1860, four 
abnormal flowers of the common Indian cress (Tropceo- 
lum ma/jus), each presenting a supernumerary spur. On 
these he remarked that "in Tropceohim 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 winch 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 addi- 
tional spurs were precisely similar to the normal ones, 
except that they were a little shorter. This abnor- 
mality, 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 coin- 
cides. The number of parts is also frequently in- 
creased ; 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 
purpwrea, in which plant prolification often adds in- 
creased 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 camjmnu- 
latum, for instance, is very generally regular. In Calceo- 
laria it is the central terminal flower which is usually 
peloriated; on the other hand, in Linaria and Antivr- 


hirwm 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 down- 
wards 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 com- 
pression of the lateral bracts is the cause of the irregu- 
larity of the andrcecium and of the receptacle. 1 

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 charac- 
terised by their deficiencies in this respect. On this 
point M. de Melicoq 2 says, referring to Linaria vul- 
garis 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 fohage, the peloriated flowers were princi- 
pally 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 multiplica- 
tion by division of the root, but Willdenow obtained 

1 Godron, " Mem. sur les Fumaviees a fl. irreg.," 'Ann. Sc. Nat.,' ser. 
5, vol. ii, tab. xvii, p. 280. 

3 'Bull. Soc. Bot. France,' vol. v, 1858, p. 701. 


seeds from the hma/ria which reproduced the 
anomaly when sown in rich soil. Baron Melicoq 
obtained similar results. 1 Mr. Darwin 3 raised sixteen 
seedling plants of a peloric Anti/rrhirmm, artificially fer- 
tilised 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 remem- 
bered, however, that in the latter cases the stamens 
are often deficient. In the Composite?, 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 com- 
munication of some flowers of Corydalis tuber osa (figs. 
124, 125), provided with two spurs of nearly equal size. 
To these flowers allusion is made in the work already 
quoted 3 in the following terms : — " Corydalis tuberosa, 
properly has one of its two nectaries colourless, desti- 
tute of nectar, only half the size of the other, and 

1 ' Bull. Soc. Bot. France,' vol. vi, 1859, p. 717. 
8 ' Variation of Anim. and Plants,' ii, p. 70. 
3 Loc. cit., p. 59. 



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 

Fig. 124. — Two-spurred flowers of Gorydalis. 

Fig. 125. — Section through two-spurred flowers of Corydalis. Magnified. 

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 Diehjtra, 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, accom- 
panied 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 Diehjtra and its allies." 

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 repre- 
sent t lie ordinary condition of another ; thus, the peloric 
Calceolarias resemble the flowers of Fabiana, and De 
Candolle, 1 comparing the peloric flowers of Swopfada- 
riacece with those of Solimacece, concluded that the 
former natural order was only an habitual alteration 
from the type of the latter. Peloric flowers of Papi- 
Uonacece in this way are indistinguishable from those 
of RosacecB. In like manner we may trace an analogy 
between the normal one-spurred Delphim/km and the 
five-spurred columbine (Aquilegia), an analogy 
strengthened by such a case as that of the five- 
spurred flower of Delphmwm 'latum, described by 
Godron. 2 The Oorydalis, 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 organ- 
ized 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 peloria3, solo sterili plantata?, degenerant 
in Linariarn," (' Sp. Plant.,' iii, p. 254); but this opinion 
is counterbalanced by thait of others, while the frequent 

1 'Theor. Elem..' ed. 2, p. 266. 

2 Cited in ■ Bull. Soc. Bot. France,' vol. xiii iRev. Bibl.). p. 81. 



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 : 

Aconituin Napellus. 
Delphinium elatuni ! 
CorydaHs tuberosa. 

* Viola odorata ! 

Impatiens Balsamina. 
Clitoria Ternatea. 
Cytisus Laburnum ! 
Trifolium repens ! 
Lupinus polypbyllus ! 
*Gloxinia, var. cult. ! 
*Linaria vulgaris ! 








cymbalaria ! 

purpiirea ! 



Digitalis orient alis. 

* purpurea ! 
Calceolaria crenatiflora. 


* var. cult. ! 
Cbelone barbata. 

* Antirrhinum majus ! 
Rhinanthus crista galli. 
Pedicularis sylvatica. 

Pedicularis eiiphrasioides. 

Scrophularia aquatica ! 

Sesamum indicum. 




Nepeta diffusa. 

Galeopsis Ladanum. 

Galeobdolon luteum. 
Teucrium campanulatum ! 
Plectranthus fruticosus. 
Cleonia lusitanica. 
Dracocephalum austrkicum. 
Phlomis f ruticosa ! 
Vitex incisa. 
Aristolochia, sp. ! 
Ophrys aranifera ! 
Orchis simia. 

pyramidalis ! 

latifolia ! 

morio ! 




Habenaria bifolia. 
Corallorhiza innata. 
Aceras anthropophora. 
Cattleya Mossia? ! 
Phalsenopsis equestris ! 
Pogonia ophioglossoides ! 

The literature of peloria is very extensive. The 
following are the principal papers, not already men- 
tioned, which relate to the subject, arranged under the 
genera, placing those first which are most subject to 
this anomaly (see also Regular Peloria). 

Irinaria. — Adanson, 'Fain. Plant.,' t. i, p. 110. Jussieu, ' Gen. Plant.,' 
p. 120. Poiret, ' Encycl. Method. Suppl.,' t. iii, Jaeger, * Missbilld. 
der Gewachs.,' pp. 94, 07, and 313. Cassini, ' Op. Phytol.,' t. ii. \>. 33] 
Ratzebourg, ' Auimadv. ad pelor. spectand.,' L825. Turpiu. ' [c. Veget.,' 


tab. xx, f. 16. Curtis, ' Flor. Londin.,' i, 118. Hopkirk, ' Flora Anom.,' 
pi. vii, figs. 1, 2. 3. galler, • Act. Helvet.,' 2, p. 25, t. iv. De Candolle, 
• Flore Franc.,' t. iii, p. 583! bowerby, 'En-l. BoO iv, 260, ed, Syme, 

tab. !'!>:>. Chavannes, 'Mon. Antirrhin.' JDelavaud, 'Bull. Soc. But. 
France,' 1S5S, p. 6X9; id., 1 Slit t, p. 17f). ] [culler, ' Linn;ea,' xvii, tab. ii. 
Webei^JVerbaiidl. des Nat. Hist. Vcrcins. f. d. Rh. Preuss.7 IS50, tab. i, 
figaTT^ST 'Verb. Nat. Hist. Ver. Rb. Prcus.,' 1819, vol. vi. p. 590, 
tab. xiii. Antirrhinum, Clos, ' Mem. Acad. TouloUB.,' \ i. L862. Clia- 
va uucs, 'Mon. Antirrb.,' p. 62. Fresenius, ' Mus. Senkcnb.,' ii, t. iv, 
fig. 10. ' Bot. Soc. Edinb.,' 1851, July 10. — Calceolaria, Cbamisso, 
' Linna?a,' t. vii, p. 200. Guilleniin, ' Arcbiv. Bot.,' t. ii, p. 1 et 136. 
Scblecbtendal, ' Linnsea,' xii, p. 686. Ernst Meyer, ' Linna3a,' xvi, 26, 
tab. iii. Morren, ' Bull. Acad. Belg.,' t. xv, n. 7, et t. xviii, p. 583. 
•Gard. Cbron.,' 1850, p. 389; ibid., 1866, p. 612.— Viola, Leers, 'Flor. 
Herborn.,' p. 145. Dc Candolle, ' Organ. Yeget.,' t. i, p. 519, pi. xlv. 
Forbes, ' Proc. Linn. Soc.,' June 6, 1848, p. 382. Hildebrand, ' Bot. 
Zeit.,' 1862, vol. xx, tab. viii. — Orchidacece, His, ' Jourl. Pbys.,' 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 Londi- 
nensis,' 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. Konigsberg,' 1860, i, 59. Masters, ' Jourl. 
Linn. Soc.,' vol. viii, p. 208 (Ophrys, Pogonia). Ducbartre, ' Bull. Soc. 
Bot. Fr.,' vol. vii, 1860, r>. 26, Cattleya. Cramer, ' Bildungsabweicb.' — 
Limosella, Baillon, ' Adansonia,' i, p. 305. (Flower normally irregular, 
becoming regular "a force d'irregularite.") — Chelone, Chamisso, 
' Linnroa,' 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 wbich 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. Yeget.,' p. 186. 
Hallier, ' Phytopathol.,' p. 151. 



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 organo- 
genists. 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 sub- 
stitute 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 pur- 
sues 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 devia- 
tions arising from similar causes will be mentioned 


elsewhere, but, under the presenl 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 teratolo- 
gical 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. 1 



This condition, wherein true leaves are substituted 
for some other organs," 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 

1 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. 

2 Engelmann makes use of the word frondescence in the same cases. 
' De Anthol.,' p. 32, § 38. while Morren adopts the term Phyllomorphy, 
' Lobelia.' p. 95. 



place of other organs, is frequently looked on as due 
to retrograde metamorphosis, or to an arrest of develop- 
ment. 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. 

Phyllody of the bracts. — As bracts are very generally 
imperfect organs, so their replacement by perfect leaves 

Fig. 126. — 'Rose plantain,' Plnntafin media var., spike contracted; 
bracts leafy. 

OF Tin: BE \> I - 


is not attributable bo arrest of development or retro- 
grade metamorphosis, but the reverse. The braetsol' 
some species of Plantago* are very subject to this change. 
Thus, in the rose plantain of gardens, /'. media (fig. 

L26), 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 /'. major (fig. 127) and in P. Icmceolata (see p. 108). 

Fig. 127. — Leaf-like bracts in Plantago major. 

It also occurs in the bracts of Corydalis solida, Amorpha 
fruticosa, Ajuga reptans, Partheniwm modorum, Gent- 
awn 1 1 Jacea, in the hrvolucral bracts of the dandelion, the 

1 See Schlechtendal, ' Bot. Zeit,.' vol. xv. 1857. p. 873; also Man-hand, 
• Atlans>mia." iv, p. 156. 



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. 1 

Fig. 128. — Dahlia. Scales of receptacle leafy. 

In TJmbelliferce the substitution of leaves for involucral 
bracts is not infrequent. It has been observed among 
other plants in Angelica Razouhii, Garwm carui, Daucus 
Garota 9 &c. The scales of the hop (Hunwlus Lvpulus) 
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, 3 
of some willows, 3 &c. The bracts of some Euphor- 
biacere, as E. pusilla, E. Lathyris, E. Oypa/rissias, have 
been observed to undergo a similar alteration. 4 

1 For instances of similar changes in Composites, see De Candolle, 
' Prod.,' t. vi, p. 571, Centaurea Jacea pliyllocephala. Clos, ' Ann. Sc. 
Nat.,' ser. iii, torn, xvi, 1851, p. 41. ' Science Gossip,' 1865, p. 104, &c. 

2 Kicks, ' Bull. Acad. Belg.,' t. xviii. part 2. p. 288. 

3 Weber, ' Verhandl. Nat, Hist. Vereins. f. Preuss.,' &c, 18fi0, p. 381. 
Weber, loc. cit. 

1 111. i \\.\ \. 2 \:> 

Amongst monocotyledons an analogous cha 
occurs not unfrequently, as in some commelynaceous 
plants, e.g. Trade8cantia s in Miosa, &c. 

The spathe ><[' Aru/m maculatv/m is sometimes repre- 
sented 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. 1 In Schcenus cephalotes a similar 
exaggerated development of the bracts is figured by 
Rottboell. a 

Phyllody in inflorescence of Conifers. — This demands | 
ing 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 fre- 
quently associated with a more or less foliaceous con- 
dition 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 de- 
scribed by Parlat ore in Abies Brunoniana, and examples 
may frequently be met with in the larch (La/rix europcea), 
and specially in Cryptomeria japonica.* The scales of 
the male catkins of conifers likewise occasionally 
assume the appearance of leaves ; this may be seen in 
monstrous catkins of Araucarza, as also in Podocarp 
and Oupressinew (Eichler). 

Phyllody of the calyx.— Sepals under ordinary circum- 
stances are so like leaves, that it is not wonderful that 

" Sauter, ' Mora v. But. Z.:-it..* 1831, p. 11. 

- 'Descr. et Icon. Plant.,' tab. 20. 

3 For references see p. 115 ; see also to Eichler, ' Excurs. Morpholog. 
de format, flor. Gynmosperni.,' in " Mart. Flor. Brasil," abstracted in 
English in ' Natural History Review.' April, 1S64. 



they are often replaced by those organs. 1 A singular 
instance of this has been mentioned as occurring in 
Gakile maritima, wherein the sepals were found by 
M. Fournier to be pinnatifid like the ordinary leaves 
of the plant. 2 The sepals of Banuncidacece and Rosacea?, 
for example, Eosa, Geum, are particularly liable to this 

Fig. 129. — Flower of rose, sepals replaced by five perfect leaves ; axis 
prolonged through the flower iu the fomi 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 liirsutum 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 

1 " Calyx tunc plane non differt a foliis proxime ipsi prsecedentibus." 
Wolff, 'Theor. Gener.,' §114. Linn., ' Proleps.,' § 6. Goethe, ' Ver- 
such.,' §§ 31-38. 

a < Bull. Soc. Bot. France,' vol. viii, 1861, p. 697. 

"i mi. calyx. 2 11 

sometimes carried to such as extent, thai five perfl 
bistipulate leaves occur in the place of the calyx, but 
when this is the case it usually happens that the 
pistil is abortive. 

FlG. 130. — Fuchsia, with one of the sepals leaf-like. 

Dc Candolle 1 fio-ures a curious instance wherein the 
pappus of Podospermwm lad was replaced by 

rive 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. AYigand 
has observed a similar transformation in a species of 
itranthus (Valeri '•). 

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 prim- 
rose 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 Oucurbita 
adherent to the carpels. It is also shown in the cut, 

1 : Organ. Veget..' t. i, p. 492, pi. xxxii. f. 6. 



fig. 132, borrowed from the ' Gardeners' Chronicle,' 
1859, p. 654. 

Fig. 131. — Primrose. Calyx of foliaceous segments. 

Fig. 132.— Leafy calyx of melon. 

OF THE ( .MAX. 249 

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 
eases; thus, in the vagina or sheath, there are gene- 
rally 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. 1 

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 ab- 
normally leafy. The complete leaf development shows 
itself more frequently among the monosepalous 

1 This distinction between laminar and vaginal venation is well seen 
in cases like Alussaenda, Calycophyllum, or Dipterocarjnis, where the en- 
larged 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). 



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 indi- 
vidual plants, when it would be found that phyllody of 
the calyx occurs much more often in individual gamo- 
sepalous 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. 

Sinapis arvensis. 

Brassica oleracea ! 

Peltaria alliacea. 
*Sisymbriuni officinale. 

Caryophyllacese, 1 sp. pi. 

Geranium, sp. ! 
*Fuchsia, var. hort. ! 

Epilobium hirsutum ! 

Cucurbita Pepo ! 
*Rosa, var. hort. ! 

Potentilla nepalensis. 

Fragaria sp. 

Geuni rivale. 

Amygdalus communis. 

Persica vulgaris. 

Cerasus ! 

Pyrus Malus. 

Daucus Carota. 

Atharnanta Cervaria. 
*Trifolium repens ! 

Centranthus macrosipbon. 

Tragopogon pratense. 

Scorzonera octangularis. 

Hypochaeris radicata. 
*Senecio vulgaris ! 

Podospermum laciniatum. 

Cirsium arvense. 

Carduus beteropbyllus. 

Campanula, sp. 

Convolvulus sepium. 
*Primula officinalis, var. cult ! 

Gentiana campestris. 
*Petunia violacea ! 

Lycium europamm. 

Laurus Sassafras. 

Tulipa Gesneriana. 

Convallaria maialis. 

Colcbicum autumnale ! (vire- 
scent ?) 

Consult also Turpin, ' Atlas de Goethe,' t. iv, f. 12, Lycium. Engel- 
mann, ' De Anthol.,' § 35, p. 31. This author figures pbyllodic 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, ' Reju- 
venescence, ' Ray Society's Transl. See succeeding paragraphs. 

i In this order Agrostemma Githago offers an illustration of a normally 
leafy calyx. 


Phyllody of the corolla. — The petals also are frequently 
replaced by leaves, though in many of the recorded 
instances bhe change lias been one of colour only; 
these latter arc strictly cases of \ hv<cence. M. Scringe 1 
Bpeaks of a flower of Peltaria alKacea 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. 

Fig. 133. — Sepals and petals to leaves. Geranium. 

The frondescent petals are very often completely dis- 
joined, as in Verbascv/m nigrum, and Lonicera Peri- 

clymemtm, in wliich, moreover, median prolification 
generally coexists. In the case of Drqpceolwn 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 Aetata 
spicata, as observed by Fresenius, the petals were 
replaced by true petiolate, palminerved, lobed leaves, 

1 'Bull. Bot.,' i. p. 6. 



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 Tropceolum 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 poly- 
petalous. 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 phyl- 
lomorphic 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 laminaB. 
(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. 

Aquilegia vulgaris. 
Actaea spicata. 
*Brassica oleracea ! 
Diplotaxis muralis. 
Hesperis niatronalis. 
Tblaspi bursa pastoris. 
Sisymbrium tenuifoliuni. 
Turritis glabra. 
Rapbanus sativus. 
Peltaria alliacea. 
Alyssum incanum. 
Erysimum Barbarea. 
officinale ! 
Cbeirantbus Cheiri. 
*Dictanmus Fraxinella ! 
Lycbnis sylvestris. 

dioica ! 
Alsine media. 
Cerastiuin vulgatum ! 

Reseda lutea. 

Malva sylvestris. 
*Trop8eolum majus ! 

Geranium, sp. ! 

Triumfetta, sp. ! 

Epilobium birsutum ! 

CEnotbera striata. 

Rubus, sp. 
*Rosa, var. cult. ! 
*Trifolium repens ! 

Spiraea oblongifolia. 

Amygdalus communis. 
*Rosa ! 

Cerasus vulgaris ! 

Persica vulgaris ! 

Potentilla nepalensis. 

Geum rivale. 

Daucus Carota ! 

Heracleum Spbondylium. 

Torilis Antbriscus. 

Echinophora maritima. 

Campanula rapunculoides. 

Pbyteuma spicatum. 

Calendula officinalis. 

Cirsiuni tricepbalodes. 

Senecio vulgaris. 

Scabiosa columbaria, 

Lonicera xylosteum. 


OF nil STAMENS. 253 

Gentians Amarella. Antirrhinum majus ! 

G-ilia glomeriflora. Stachys sylvatica. 

•Symphytum officinale. *Aiiagallia phceniceaP 
Petunia violacea I Primula Brner i ! 

Verbascum, sp. Poleinoniuin cceruleuni. 

See Moquin-Tandon, 'El. Terat. Veg.,' p. 203. Engelmann, ' De 

Anthol.,' § '38 et seq. ; tab. ii, figs. 8 — II. Quia ; bab. v. ■_;:!— 2<>, Benecio . 
tab. v, f. 1—13, TaHMs ; tab. iv, f. 3, Erysimum. ' Bull. Soc. Bot. Fr.,' 
vol. ii, 1855, p. 479, Primula sinensis. Giraud, ' Edinb. Phil. Magazine,' 
1839, Antirrhi tin hi. Jaeger, 'Act. Acad. Caes. Nat. Cur.,' vol. xiii, 2, 
p. 1, tab. xli, Tropceolum. Bischoff, ' Lehrbuch,' 11, 2, p. 27, note, 
TropcBolum. Fresenius, ' Mus. Senkeub.,' ii, 35, tab. 4, fig. 5, AcUca. See 
also succeeding pai-agraphs and sections in Chloranthy, Yirescence, &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 develop- 
ment than that which occurs in those cases where 
less highly differentiated organs, such as the sepals, 
petals, and pistils, are thus altered. 1 

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, repre- 
senting 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 inflor- 
escence, some of the stamens were partly frondescent, 

1 Wolff's original opinion was tbat tbe stamens were equivalent to so 
many buds placed in tbe axil of tbe petals or sepals (see ' Theoria Gene- 
rationis,' 1759, § 114) — an opinion wbicb more recently bas received tbe 
support of Agardb and Endlicber. Wolff himself, however, seems to 
have abandoned bis original notion, for in his memoir, " De formatione 
intestinorum pracipue turn et de amnio spurio aliisque partibus embry- 
onis gallinacei, nondum visis," &c, in ' Comm. Acad. Petrop.,' xii, p. 403, 
anno 1766, be considers the stamens as essentially leaves. See also 
Linn. 'Prolepsis,' § viii ; Goethe. ' Metam..' § 46. 



half the anther being perfect, the other half leaf-like. 
Another filament bore just above the usual joint 
three leaflets, two lateral ones, somewhat con- 

Fig. 134.— Flower of a Petunia, opened to show the stamens partially 
replaced by stalked leaves. 

duplicate, and a third central one, half anther, half 

In the case of frondescent flowers of Tropceolu/m 
ma jus 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-lobecl leaf, the terminal lobe represented by 
the anther. 

In Jatropha Pohliwna, Mull. (Admorophwm hum- 
rians, Pohl.), a singular condition has been observed by 
M. Midler (Argov.). In this flower the anther, in place 



of being representee! 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. 130), thus resem- 
bling the cases of adhesion of leaves by their Burf 
already referred to (p. 33). In other cases, in the same 
plant, the anther appeared as if formed by two collateral 

Fig. 135.— Phylloid anther of 
Jatropha, after Miiller (Arg.). 

Fig. 136.— Leaf-like anther of 
Jatropha Polrfiana. after Miiller. 

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. 1 The bear- 
ings of these and other similar malformations on the 
morphology of the anther are alluded to under the 
head of petalody of the anther. 

Pliyllody of the stamens has been most often observed 
in the following plants : 

1 Miiller (Argov.), in ' Mem. Soc. Phys. et d'Hist. Xat. Genev..' t. xvii. 


Anemone nemorosa. Daucu3 Carota. 

coronaria. Epilobium hirsutum ! 

Delphinium crassicaule. *Rosa, var. cult. ! 

Nymphsea dentata. Lonicera Periclymenum. 

Tropseolum majus ! Anagallis arvensis. 

Dictamnus albus. Primula sinensis ! 

*Trifolium repensi Petunia, var. cult. 

Torilis anthriscus. Jatropha Pohliana. 

Heracleum Sphondylium. Euphorbia geniculata. 

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. 1 — This is of more common occur- 
rence 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 lamina?, 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 3 
relates having found in the neighbourhood of Mont- 
pellier a flower of a tulip the ovary of which was repre- 
sented by true leaves, which bore on then margins the 
ovules, and thus presented a striking analogy with 
the carpels of those Sterculias, like 8. platcmifolia, 
which are foliaceous in texture and open very early in 
the course of their development. A similar occur - 

1 "• 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 combi- 
nations." — (Goethe, ' Metam.,' § 78.) Wolff, ' N. Comm. Acad. Petrop.,' 
1766, xii, p. 403, expresses precisely the same opinion as to the nature 
of the seed-vessel. 

2 'El. Terat. Veg.,' p. 205. 



rencc lias also been frequently noticed in the Columbine 
and also in Grucif eras and Umbelliferce. M. Germain de 
St. Pierre mentions an instance wherein the carpels of 
Saiix Babylonica were converted into two leaves, pro- 
vided with stipules. All the flowers of the catkins 

Fig. 137. — Rose, in which the axial portion <>f the flower was elon- 
gated and the carpels were more or less replaced by leaves. 




were similarly changed, so that it became permanent, 
and resembled a branch. 

Substitutions of this kind form the green " eyes" or 
centres of certain varieties of Banunculus 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. 

Fio. 138.— Cucumber with leaf attached. 


Pig. L38, for which I am indebted to Mr. S. J, 
Salter, represents a Nrery singular conformation in the 
cucumber, described by thai gentleman in 'Henfr< 
Botanical Gazette,' i, p. 208, and considered by him to 
be due fco 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 attend- 
ing the acceptance of this explanation reside in the 
peculiar reversed position of the leaf, and in the fact 
that the fruit of the GucurbitcLcece 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 1 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 congeni- 
tally 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 con- 

1 -Ann. Sc. Nat., 5 ith series, v..l. be, p. 209. 



111 the preceding instances the foliaceous condition 
has pervaded the entire pistil, or at any rate the basal 

Fig. 139. — Flower of Trmmfetta, sp., carpels represented by five 

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 CerasuSy Daucus, &c. 

There are cases, however, in which a part only 
of the pistillary structure thus becomes foliaceous. 
Linnaeus, ' Prolepsis,' § 9, mentions some flowers of 
Carduus Ueterophyllm and G. 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, 1 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 vagini- 
form part of the leaf; the three styles were formed by 
the petioles, while the stigmas were represented by 
tri-foliolate leaves. The back of the leaf in these 
cases is usually directed away from the centre of the 

1 ' Adansonia,' iv, p. 70. A similar deviation has been observed by 
M. van Tieghem in the ovary of Tropceolum magus, ' Bull. Soc. Bot. Fr.,' 
1865; p. 411. 


flow or. 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 Umbelliferce, 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 folia- 
ceous pistils, they may be placed, as in Aquilegia, 
DeVphmnm, &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. 1 The 
supposed causes of this and other similar malforma- 
tions 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 frond- 
escence of the pistils. 

As this condition rarely occurs without correspond- 
ing changes in other parts of the flower, further re- 
marks 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 : 

Pceonia officinalis. Nymphsea dentata. 

Ranunculus repens ! Sinapis arvensis ! 

*Aquilegia vulgaris ! Diplotaxis tenuifolia. 

Delphinium elatum. *Brassica olei'acea ! 

crassicaule. *Sisymbrium officinale ! 

Ajacis. Dianthus, sp. 

ainsenum. Reseda Phyteuma. 

1 Planchon et Mares, ' Ann. Sc. Nat.,' ser. 5, vol. vi, 1866 ; p. 228, tab. 


Triumfetta, sp. ! Melilotus, sp. 

Lyclinis dioica. Medicago, sp. 

Oerastium, sp. ! Lonicera Periclymenum. 

*Dictainnus Fraxinella ! Carduus heterophyllus. 

Oerasus avium. tataricus. 

vulgaris ! Scrophulavia aquatica. 

*Rosa, var. cult. ! ■ Symphytum officinale. 

*Daucus Carota ! Anctusa ochroleuca. 

Heracleum, sp. paniculata. 

Ej)ilobium liirsutum ! *Primula sinensis ! 

Lathyrus latifolius. Salix babylonica. 

*Trifolium repens ! Hyacinthus, sp. 

hybridum. Tulipa, sp. 

Some of tlie above are probably cases of mere 
virescence rather than of pliyllody. For further illus- 
trations, 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 evi- 
dence is at present very conflicting. Prof. Al. Braun, 
who has devoted much attention to the subject, de- 
scribes 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 pro- 
nounce an opinion on the nature of the pedicel of the 
ovule. In PrimulacecB, 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 

01' THE 0\ ULES. -<' :> > 

ovules of Prirrmla sinensis, applied the term Lepyro- 
I >I i \ H \ (Xerrvpbv, a scale) to the foliaceous condition of 
the testa in this plant. Qnger 1 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 lias been de- 
scribed by A. de Candolle. 3 

In these flowers the placenta seemed to be composed 
of several funiculi soldered together, and bearing imper- 
fect 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 
Gortusa Mathioli, in which the funiculi bore little 
rounded leaves. Brongniart has described some mal- 
formations of Primula sinensis in which the ovules 
were transformed wholly or partially into small leaves 
with three to five lobes. 3 Dr. Marchand 1 mentions 
similar changes in Anagallis <irrcnsisix\\c\Lonicera Peri- 

Cramer 5 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. Tas si 6 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 

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 

1 ' Act. Acad. Nat. Cur.,' 22, 11, 1850, p. 513, t. v, vi. 

2 ' Nene Denkschrift der atlg. Scliweiz. Gesellscli.,' band v, p. 9, tab. 3, -1. 

3 ' Aim. Sc. Nat.,' 2 ser., vol. i, p. 308, pi. ix, c. 
* ' Adansonia,' vol. iv, pp. 159, 171. 

•■ • Bildungsabweickungen,' &c, tab. iv, figs. 1, 2, 21, 28, 29. &c 
6 'Bull. Soc. Bot, France,' viii, p. 395. 



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. 

Fig. 140. — Sinapis, replum and ovules ; the dotted line shows the 
position of the carpels. 

Griffith, in alluding to a similar case in Sinapis, 1 
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 origi- 
nate as a little bud from the surface of the leafy 
ovule (figs. 141, 142). 

1 ' Notulse,' p. 125, atlas, pi. xxxv ; and ' Journals of Travels,' 1847, 
p. 475, Lonicera. 

t»F TIIK <>\ UI.KS. 


In :i species of Trmmfetta (seep. 2G0), of which I 
examined dried specimens, the 
ovary was open and partly folia- 
ceous ; it bore on its infolded 
margins ten erect leaflets, repre- 
senting so many ovules ; eacli 
leaflet was conduplicate, the back 
being turned towards the pla- 

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 carpel- 
lary leaf itself — to be, as it were, 
a lobule or small process of the 
carpel, and not an absolutely 
new growth. Thus, Planchon, 1 
from an examination of some 
monstrous flowers of Drosera in- 
termedia, was led to the inference 
that the ovules are analogous to &c, Trifolivm repens. 
hairs on the margins of the 
leaves. This acute botanist was 

Fig. 141. — Leafy ovules, 

Fig. 142. — Leafy ovules of Trifolium repens, showing formation of 
nucleus, &c. After Caspary. 

enabled to trace all the gradations between the simple 

1 ' Ann. Science Nat.,' 3rd ser., vol. ix, p. 86, tabs. 5, 6. 



cup formed by the confluence of four glanduliferous 
hairs and the concave leaf and the perfect ovule. 

Brongniart 1 records ovules of Delphinium elatum 
existing in the form of marginal lobes of the car- 
pellary 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. Olos 3 mentions a similar in- 
stance in Aquilegia Shinneri ; and another is figured in 
Lindley's ' Elements of Botany,' p. 88, f. 180. 

Cramer, 3 from an examination of several ovular mal- 
formations, 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 

Eig. 143.— Portion of an open foliaceous carpel of Delphinium, with 
ovules on the lobules. 

1 ' Comptes Rendus/ vol. xviii, March 25th, 1864, and ' Ann. Sc. Nat.,' 
3 ser., vol. ii, p. 32. 

2 ' Mem. Acad. Sc. Toulous.,' ser. 5, vol. iii. 

3 ' Bildungsabweich. Pflanz. Famil.,' p. 8!>. tab. xi. 

OK Till. "\ i LES. 


from the modified lobes of the carpellary Leaf. Figs. 
143 — 145, copied from Cramer, Bhow how the nucleus 

of the ovule is formed as a new growth from the 
surface of the lobes of the leaf in Delphinium elatum. 

Fig. 1J4. — Section through marginal lobe of carpel (Delphinium), show- 
ing the nucleus (n). 

Fig. 145. — Section through marginal lobe of carpel, showing nucleus 
and tegument (DelpMniu/m). 

One of the most singular instances of ovular mal- 
formation on record is that cited by the Rev. M. J. 
Berkeley, in the ' Gardeners' 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. 146. — 1. Placenta of Diantlms, bearing ovules and carpels. 
2. One of the ovaries separated. 

FlG. 147. — Ovules of Diauthus passing into carpels. 

(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 sec- 
tions at e, (/, e. Could such a change occur in the 
animal kingdom, there would be the unfertilised ovum 

OF 1 UK >»\ I LES. 269 

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 Mas 
not developed, and sufficient evidence has been above 
given as to the foliar nature of the priniine, while for a 
leaf to be folded up so as to form a carpel is an ordi- 
nary occurrence. 

It is worthy of remark that in these foiiaceous 
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 foiiaceous condition in 
the carpel, the margins of which are more or less dis- 

Prof. A. Braun remarks that up to this date no such 
change has been observed in the ovules of Monocoty- 

Changes in the nucleus of the ovule. — The preceding re- 
marks 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 indi- 
cations of anthers. Wycller has observed a similar 
occurrence in ovules ofAlliaria officinalis, and Schimper 
has described and figured specimens of Nig ell a dama- 
scened 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 Graillardia, 
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 develop- 
ment of the nucleus, or whether it was wholly inde- 
pendent of the ovule. 

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 conclu- 
sions from the foliaceous ovules of Trifoliwn 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 integu- 
ment. The nucleus itself is considered to be a new 
formation analogous to a shoot. 



M. van Tieghem's conclusion 1 from the examination 
of flowers of Vropceolu/m majus, in which the ovules 
were replaced by perfecl peltate Leaves, is that the 
ovules are foliar productions springing, not directly 
from a prolonged floral axis, as in Prvm/ulacece, 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 i 

*Aquilegia vulgaris ! 

Delphinium crassicaule. 

Nigella damascena. 

Adonis autnmnalis. 

Cheiranthus Cheiri ! 

Nasturtium, sp. 

Sisymbrium officinale ! 

Brassica napus ! 
* oleracea ! 

*Alliaria officinalis ! 

Sinapis arvensis ! 

Turritis, sp. 

Thlaspi arvense. 

Erucastrurn Pollicbii. 

Stellaria media. 
*Reseda lutea. 

Drosera intermedia. 

Agrosteruma Githago. 

Stellaria media. 

Triumfetta, sp. ! 

Tropseolum majus ! 

Dictamnus albus. 

Fraxinella ! 

Carum carni. 

Pastinaca sativa. 
Torilis anthriscus. 
Thyssulinurn palustiv. 
Epilobiam padustre. 
Rosa. sp. 
Fragaria alpina. 
; TrilY4ium repens ! 
Medicago maculata. 
Desmodium canadense. 
Melilotus macrorhiza. 
Lonicera, sp. 
Gaillardia ! 
Crepis, sp. 
Pbyteuma odorata. 
Symphytum Zeyberi. 

Stachys sylvatica. 
AnagalLis arvensis. 
Lysimacnia ephemenim. 
*Primula sinensis ! 
Gilia glomeruliflora. 
Riimex arifoHus. 
Salix caprsea. 

The following list of publications relating to ovular 
malformations is copied from A. Braun, ' Ueber Polyem- 
bryonie und Keimung von Caelobogyne' (Appendix), 2 to 
which are also added some others not alluded to by 

1 'Bull. Soc. Bot. Fr.,' 1865. p. 411. 

" Translate! in -Ann. So. Nat., 5 4th series, t. xiv. p. 24. 


that author and not specially referred to in the pre- 
ceding pages : 

Jaeger, 'Missbilld. d. Gewachse,' p. 78, 79, f. 47. Beeper, ' Enuin. 
Eupliorb./ 1824, p. 45, Delphinium. — Schimper, 'Flora,' 1829, pp. 437-8, 
et ' Mag. fur Pharinacie de Geiger,' 1829-30, pi. iv — vi, text wanting, 
Primula, Beseda, Cheirardhus. — Engelmann, 'De Antholjsi,' 1832. — 
Valentin, ' Act. Acad. Xat. Cur.,' 1839, p. 225. Lytimaeltia. — 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, pi. iv, v, Primula. — 
Reissek, ' Linna-a.' xvii. 1843. AMaria. — Wydler, ' Denkshrift. d. Begensb. 
Bot. Gesell.,' 1855, iv, s. 77, t. vii, Alliaria.- — "Wigand. ' Grundlegung 
der Pflanzen Tevatol.,' 1850, p. 39. TurriMs.— Wigand, 'Bot. Fntersuch- 
ungen,' 1853, p. 23, Rosa, Turritis, Grepis.— Germain de St. Pierre, 
' L'Institut,' 1853, n. 1051, p. 351. — Bossmann, " Entwicklung der 
Eiknospen aus deni Fruclitblatte," <fcc, ' 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. .' . — Springe and Heyland. 'Bull. Bot.,' 1 — 7, 

Dvplotaads. — Clos, 'Mem. Acad. Toulouse.' vi. 1^62. Delpihinium. — Morren, 
C., 'Bull. Acad. Belg.,' xis, part ii, p. 519, Primula. — Caspary, 'Sclrrift. 
d. Pkysik. (Ek. Gesell. zu Konigsberg,' band ii, p. 51, tabs, ii, iii. 
Fleischer, ' Ueber Missbildnngen Versekiedener Cultur Pflanzen.,' &c., 
Esslingen, 1862. Cramer, ' Bildungsabweich,' p. 68, See. &c., Trifolium. — 
Moquin-Tandon, ' El. Terat. Veg.,' p. 206, Cortusa. — Guillard, ' Bull. 
Soc. Bot. Fr.,' 1857, vol. iv, p. 761, Stellaria. — Moelkenboer, ' Tijdschrif t 
v. Xatuurl. Geschied.,' 1843, p. 355, t. vi. vii, Primula. — Van Tieghem, 
'Bull. Soc. Bot. Fr.,' 1865, p. 411, TropcBolwn. 

Phyllody in accessory organs. — In addition to the ordi- 
nary organs of the plant, what are termed the acces- 
sory 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 occa- 
sionally in Berberis; a similar thing occurs in the 
stipules of some Legvmmosce ; the scales of some 
begonias; the tendrils of Big'i . ' ; i . Are. 

The presence of two small green laminae on the 
outer side of the two posterior stamens in Antirrhirwm 
majus has also been met with. The adventitious 
organs appeared as if they wire developments from 
the thalamus — a kind of foliaceous disc, in fact. 


Chloranthy. — r JMic term phyllomorphy is applied fco the 

individual pails of the flower wliidi assume the form 
and appearance of leaves. By chloranthy il is to be 
understood thai all, or the great majority of the organs 
of the flower assume these conditions. 1 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. 

Fig. 149.— Leafy petal of Epi- Fig. 150.— Chloranthy, &c. Epi 

lobhim. lobium liirsutvm. 

Chloranthy, then, is a more complete form of frondes- 
cence. 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 meanl 
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 

1 The calyx is not unfreqnently excepted. 




the instances to be now mentioned were not strictly 
cases of chloranthy. 

Seringe 1 has described a malformation in Diphtaads 
t&n/wifolia in which all the floral organs were replaced 
by sixteen distinct leaflets which had preserved their 
proper relative position. The Cruciferce, of which 
family the last-named plant is a member, are particu- 
larly liable to this malformation, as also are the 
Bosacece, 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 arose 2 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 

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 ruicropyle. 

1 ' Bull. Bot.,' t. i, p. 6. 

- Lindley, 'Theor. Horticult,,' ed. 2, p. 84. f. 17. 


becomes in process oftime suppress) d. Get m coccineum 
has been found byWigand with its flowers in this con- 

Lindley* figures a very interesting illustration in 
Potentilla nepalensis, in which some of the flow 
have their component parts Leafy, in others the recep- 
tacle 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 xUpine 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. 1 

1 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. 1- 

a ' Flora,' 1856. p. 711. 

s ' Theory of Horticult.,' ed. 2. p. 90, f. 25. 

4 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 {Frag fa hiapido) is a sort of 

botanical Dodo upon which many have written, and which few have 
seen. Many years have elapsed since it was first di- and 

although a century and a half have passed since there was any evid 
of its existence, it serves still as an illustration for students in mor- 
phology 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 i - 
preserved." Doubtless one of those " lovers*' was his friend John Parkin- 
son, 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 ordi- 
nary, 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 
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 straw- 
berry, but with many small harmlesse 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 

27G niYLLODY. 

Among Leguminosce a partial leafy condition (frond- 
escence), 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, 1 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 com- 
paratively 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 papilio- 
naceous 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, gi-een, 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, 

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 : " Canta- 
brigian 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 Philosophic 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. II paroit ne pas avoir aujourd'hui 
plus d'existence." 

i 'Bull. Soc. Bot. France,' 1856, vol. iii. p. 477. 


the corolla being then absent. The heads of flowers in this first form 
have the aspect of little tofts of leaves. 

'2. Each of the teeth of the calyx is represented by a long stalk, termi- 
nated by a single articulated leaflet, the bi-labiate form of the calyx is 
still recognisable; the two upper petals are united, .the three lower 
eparate; 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 papilionaceotis, the filaments free, the car- 
pellary 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 foliaceons, 
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 f oliaceous condition. Inflor- 
escence — 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, 
an'ested 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 18-49 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 Nymphcea Lotus have been 
seen in which all the parts of the flower, even to the 
stigmas, were leafy, while the ovules were entirely 

Planchon 1 figures and describes a flower of Brosera 
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 sti- 
pules, and were circinnate in vernation. 

1 'Ann. Se. Nat.,' 3 ser., vol. i\. p. sti. tabs. v. vi. 


M. A. Viand- Grand-Marais 1 records an interesting 
example of chloranthy, in which the sepals, petals, 
pistils, and ovules of Anagallis arvensis were all folia- 
ceous. Similar changes have not unfrequently been 
met with m'Dictamiius Fraxinella. 

M. Germain de Saint Pierre has also recorded the 
following deviations in the flowers of Rumex arifolius 
and B. scuta t us ; in these specimens the calyx was 
normal, the petals large, foliaceous, shaped hke 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 folia- 
ceous appendage analogous to the primine. 2 

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, 3 consisted below of the ordinary unchanged 
glumes, but the remaining palese as well as the lodicles 
and stamens were represented by ligulate leaA*es. The 
plant, it is stated, was affected by a parasitic fungus. 
On the other hand, General Munro, in his valuable 
monograph of the Bambiisacecef 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, how- 
ever, perfect fertile spiculae; as progress is made to- 
wards 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 

1 ' BuU. Soc. Bot. France,' vol. viii. 1861, p. 695. 

- Ibid., vol. iii, 1856, p. I ' 

* 'Flora/ 1856, p. 712. 

4 ' Trans. Linn. Soc.,' vol xxvi, p. 37. 


remarks with justice that the position of the flowers 
on the axis is ol* 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 J&vibus there are two sorts of chloranthy, ac- 
cording 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 inflor- 
escence, very loose and indeterminate, and with 
axillary flowers. 1 

On the whole, taking in consideration cases of par- 
tial 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 2 gives an instance 
in Stella/ria media where the condition appeared to be 
due to the attacks of an insect Tlirvps 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. 

1 'Bull. Soc. Bot. France,' 18G2. vol. ix, p. 3G, tab. i, and also p. 291. 

2 Had.. 1857, vol. iv, p. 701. 

280 PHYLLOM". 

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 deve- 
lopment 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 refer- 
ence should also be made to preceding and subsequent 
sections, and to that relating to prolification of the 

Aquilegia vulgaris. Selinum caruifulium. 

Chelidoniuin majus. Epilobiuin hirsutum ! 

Corydalis aurea. Begonia fuchsioides. 

Xymphaea Lotus ! Gomphia, sp. 

*Brassica oleracea ! Scabiosa Columbaria. 

Bunias. Dipsacus fullonum. 

Hesperis matronalis. Matricaria Partheniuin. 

*Sinapis arvensis ! Calendula officinalis. 

Sisymbrium officinale. Campanula pyrainidalis. 

Erucastrum canariense. Reseda odorata ! 

Diplotaxis tenuifolia. Vitis vinifera. 

Lychnis dioica ! Dictamnus Fraxinella ! 

Cerastium glomeratum ! Triumfetta, sp. ! 

triviale. *Trop8Bolum majus! 

Stellaria media. Rhamnns Frangula. 

Poterium polygamum. *Trifoliuni repens ! 

Torilis antbriscus. Lupinus, sp. 

Se3eli, sp. Rosa diversifolia ! 


Potent ilia nepalensis. AnagallxB arvensis. 

argentea. Webbiana. 

Fragaria vesc i I Nicotiana rnstica. 

Geuin rival'. Anchusa ochroleuca. 

Rubus fruticosus. Myosotis csespitosa. 
caesius. liys sylvatica. 

ifraga foliosa. Gilia capitata. 

Verbascum phlomoides. Euphorbia Begi talis. 

Scrupbulai-ia nodosa. Rumex arifolius. 
acroatica ! scutatus. 

*Primula sinensis ! Juncus lanipocarpus. 
Lysimachia Epheineruin. uliginosus. 

In addition to the publications before cited the fol- 
lowing may be named as containing valuable informa- 
tion on the subject of this chapter. 

Jaeger, ' Missbild. Gewiichs.,' 181-4, p. 83, Trifoliwm repens. For other 
accounts of similar malformations in tbe same plant, see Scbmitz, 
" Linna?a,' xv, p. 268. Unger, ' Flora' (B. Z.) xxv, p. 369. Caspary, 
' Schrift. der. Physik. okon. Gesellsch. zu Konigsberg,' 2, 1861, p. 51, 
tabs, ii, iii. Fleischer, ' Missbilld. verschied. Cult. Pflanz.,' 1862, p. 56, 
&c.j 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.' 
MoiTen, C, ' Bull. Acad. Roy. Belg.,' xix, part 2, p. 539. Molkenboer, 
' Tijdschr. voor Xatuurl. Geschied.,' 1843, p. 355, tabs, vi, vii. Marchand, 
' Adansonia,' iv, p. 167 and p. 159, AnagaUis, p. 171, Lonicera, p. 83, 
Juncus. For other plants see Presenilis, 'Mus. Senk.,' 2, p. 35, &c. 
Xorman, 'Ann. Sc. Nat., 9 ser. 4, 1858, vol. ix, p. 220. Christ, 'Flora' 
(B. Z.) 1867, p. 376, tabs, v, vi. Stachys. Cramer, ' Bildungsabweich.,' 
p. 26. Ac. Baillon, ' Adansonia,' ii, p. 300. Moquin-Tandon, ' El. Ter. 
Veg..' p. 230. Schauer's translation, p. 220. Hallier, ' Phytopathologie,' 
p. 160. 



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 fre- 
quent 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, substi- 
tution, 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 discrimina- 
tion 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 
Fig. 152.— Flower question there are neither stamens 
of St. Valery apple, nor petals, unless the second or inner 
with sepaioid pe- row f se p a ] s fo e considered as sepa- 

loid petals (fig. 152). 

M. Alph. de Candolle 1 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 

1 'Neue Denkschriffc. Schweiz. Gesellsch.,' band v, p. 9. 


the (lower to sepals bave occurred in monocotyledonous 
plants, or others in wliicli 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 seg- 
ments 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 

Petalody of the calyx— Calycanthemy. — As with the bracts, 
so the calyx in certain instances is naturally coloured, 
as in Delphinium, Tropceolwm, and others. In Mus- 
scenda, Galycophyllwm, listeria, &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 cultiva- 
tion a variety of the primrose called Primula calycan- 
thema, 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 1 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 

1 ' Bull. Acad. Belg.,' xix, part 2, p. 93. 


proper to the corolla is developed in the tube of the 

Fig. 153. — Flower of Minnulus, with petaloid calyx. 

calyx, the edges of the limb remaining green, the middle 
of the limb being purple {Primula tricolor). 

Under this head may be mentioned the occurrence of 
tubular sepals in place of the ordinary flat ones in 
Helleborus olym/picus ; only two of the sepals were thus 
affected in a specimen recently observed — a third ex- 
hibited 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 men- 
tioned, and also in the following : — Ga/m/pa/rmla persici- 
folia, Anagallis arvensis, Gloxinia, Syringa persica? 
Calceolaria, §c. Sfc. In the last-named plant one or 
more of the lobes of the calyx may frequently be seen 
replaced by a slipper-like petal. 

1 Schlt'clitendal, ' Linnsea,' ix, p. 737. 


Among polysepalous plants petaloid sepals bave 
been observed in Ra/n/imcul/us (mricorrms, Tt/ubus ccesvus, 
Sfc Fleischer also describes a case of this kind in 
( 'a/mm carui? 

It will be seen from the above thai 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 orchi- 
daceous 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) 2 
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, tri- 
plices, &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 Nyw/phcea, Carina, 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 con- 
junction with an increased development of parts (Mul- 
tiplication), or with a similar change in the carpels, 
and it is either partial or complete. 

i Misbilld., « Cult. Gewachs..' p. 32. 
2 Linn., ' Phil. Botan.,' § 120. 


Among the flowers in which petaloid development 
of the stamens happens most frequently may be men- 
tioned 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. More- 
over, 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 occu- 
pied by a series of petal-like structures entirely desti- 
tute 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 in- 
creased 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 tuhps 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 appear- 
ances have been observed in Iris, Hyacinths, Nar- 



cissus, Colchicum, and Crocus. M. Fouraier 1 describes 
a flower of Na/rcissus Tazetta from within the normal 
perianth of which sprang a second one, equally pro- 
vided will) 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. 2 

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. 3 For instance, in Sulanum 

Fig. 154. — Double columbine, Aquilegia — petalody of the filament. 

1 'Bull. Soc. Bot. France,' 1859, vol. vi, p. 199. 

2 Seeman's ' Journal of Botany,' vol. iii, p. 105 ; also Morren, ' Bull. 
Acad. Belg.,' vol. xx, part 2, p. 264. 

3 Morren, 'Bull. Belg.,' xviii, p. 503. 


tuberosum, S. Dulcamara, in Anagallis, in Fuchsia, 

and some other plants, the anther-lobes themselves 
become petaloid, while the filament remains un- 

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 1 observes that in the Rammculcicew 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 Mallow.-. 
Roses, Magnolias, &c, petaloid expansion of the fila- 
ment is most common, though it is by no means con- 
fined to such flowers, the change occurring in Alia- 
manda cathartica, Jasminum grandifiorum, 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 

In those cases where there is more than one row of 
stamens, the outermost are most liable to this change : 
thus in Samfraga decvpiens, as shown by Ch. Morren, 2 
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 mosl 

1 ' Organ. Veg.,' t. i, p. 513. 

2 ' Bull. Acad. Roy. Belg.,' tome xvii; and Lobelia, p. 65. 


flowers tliat have their stamens in two rows. Occa- 
sionally it happens that an outer scries of stamens is 
abortive, or wholly suppressed, while the inner row 
becomes petalodic ; this was the case in some flowers 
of TAlvu/m an r<i hi hi 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 pre- 
sence of four wing-like processes emanating from the 
central filaments, two on each side, so that the arrange- 
ment 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 ordi- 
nary circumstances, produces pollen. In the malformed 
flowers no pollen is formed, at least in the more com- 
plete states of the malformation, but the walls of the 
anther lobe become preternaturally enlarged, and peta- 
loid 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 per- 
fect pollen-sacs or quarter-anthers, the remaining por- 




tions being petaloid and continuous with the dilated 
filament. Not unfrequently these semi-petaloid sta- 


Fig. 155. — Four-winged filaments of Rhododendron. 

mens 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. 1 (See ante, p. 35, fig. 12.) 

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 Ptercmd/ra, 
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 Orchidacece generally arise from 
petalincation 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 sup- 
pressed, so that the morphological structure of the 
flower, at first a matter of theory, becomes actually 

1 Masters, " On Double Flowers," ' Rep. Tnternat. Bot. Congress.' 
London, 1866. p. 127. 


realised. Pig. L56 is a diagram Bhowing the presence 

of two additional labella within the ordinary one in 
a specie- of ( >atasi turn, and repre- 
senting two petaloid Btamens, 
thus evidently completing the 
outer stamina! whorl, of which 
there is usually but a single 
representative (see Peloria, Mul- 
tiplication, Prolification). In 
some of these double orchids it is, FlG i^.— Diagram of 
however, necessary not to con- flower of Catasetum, with 
found a petaloid condition of the tw ° labella - 
existing column with the development of usually sup- 
pressed stamens in a petaloid form. Thus, in Lycaste 
SJrinneri 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 suffi- 
cient to enable the observer to ascertain the true 
nature of the appearance. 1 

Some forms of duplicate or hose in hose corollas art' 
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." 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 

When the petalody specially affects the anther-lobes, 
as in Arbutus, Petunia, Fuchsia, Sfc, the venation of 
the petal-like portion is very frequently laminar, thus 

1 See also C. Morren, ' ; Stir les waies fleurs doubles chez les Orchi- 
dees," 'Bull. Acad. Roy. Belg..' vol. xix, pan; ii. 18-52. p. 171. 

J C. Morren, ' Bull. Acad. Belg.,' vol. xx. 1853. part ii. p. -2-1 

3 • Rep. Bet. Congress.' London. lSGG. p. 135. t. vii. f. 14. 


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. 1 

1 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 Iriiat 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 fila- 
ments 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 
fine 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 be- 
tween these plates corresponds thus to the suture 
of the normal anther. 

The inner or upper portion of the anther-leaf 

is that which is most intimately concerned in the 

/#'?AT"7(Vf^N formation of pollen ; it comparatively rarely 

\ y \ \ (query ever) happens that the back or lower 

■p i-n td surface of the antheral leaf is specially devoted 

J!IG. loi.—fcta- to tte f ormat i on f pollen. On the other hand, 

ioid stamen ot ^ cageg ^ Q t h ose f ^q common houseleek, 

viola, with tour w h ere we meet with petaloid organs combining 

projecting plates. the attr i Dute8 f 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 Malvacets, in 
which the stamens are frequently partly petal-like, partly divided into 



Petalody of the connective is of Less frequent occur- 
rence than the corresponding change in the other 
portions of the stamen. It may be seen in some 

forms of double columbine, 1 in which the connective 

Fig. 158. — Portion of a double columbine (AqwMegia), showing petalody 
of the connective. 

forms a tubular petal or nectary, and in double 
petunias and fuchsias. When it occurs, the true 

numerous separate filaments, each bearing a one-, or it may be even a 
two-lobed anther. This circumstance is confirmatory of the opinion 

Fig. 159. — Petaloid stamens, Hibiscus. 

held by Payer, Duchartre, Dickson, and other organogenists, as to the 
compound nature of the stamens in these plants. The stamens are 
here the 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. Midler's paper on the anther of 
Jatropha Pohliana, &c, referred to at page 255. 

1 See C. Morren, "On Spur-shaped Nectarines," &€., 'Ann. Nat. 
Eist.,' March, 1841. p. 1, tab. 11. 


anther-lobes are usually atrophied, and little or no 
pollen is formed. 

An occurrence of this nature in Tacsonia pvrmati- 
stvpula, in conjunction with the partial detachment of 
the stamens from the gynophore, led Karsten to estab- 
lish a genus which he called Poggenclorffia} 

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 (Polypetala3, 
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. 2 • The presence of these compound sta- 

1 Karsten, ' Flor. Columb. Spec.,' tab. xxix. 

2 See Dickson, " On Diplostemonous Flowers," ' Trans. Bot. Soc. 
Edin.,' vol. viii, p. 100; and on the Andrceciuni of Mentzelia, &c, in 
Seemann's ' Journal of Botany,' vol. iii, p. 209, and vol. iv (1866), p. 273 
(Potentilla, iCr.). 

I - 1 - 1 \I.I>I'\ 

mens affords a satisfactory explanation of the appear- 
ance in some double Malvaceae, wherein the tufts of 
adventitious petals arc very liable to be mistakerj 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 ! — 2. 
/ *Clematis ! 
/ *Hepatica ! 
/ Ficaria! 
/ Tbalictrum. 
/ Oaltha! 
/ »Trollius ! 
Nigella, : 
/ *Aqtdlegia ! 
/ ^Delphinium ! 
/ *Adonis! 







*Paeonia ! 
*Xelunibium ! 
*Nyinphaea ! 
*Berberis ! 
^ PapavcH — 
*Glielidoniu m ! 
*Mathiola ! 
*Gheirantlius ! 
is ! 

lamine '. 
*Hesperis , 
*Barbarea ! 
(_, *Brassica ! 
-T^Heliantkeniuin ! 

<f *Diantbus ! 
f *Saponaria ! 
i *Lychnia ! 
f *Silene ! 
; ina ! 

haea ! 
dva : 

/ 2~ iEsculus ! 
/ 2 > *Geranium ! 
/ ^Pelargonium 
XiJ. *Tropacolum .' 

T /j/Oxalis! 
/<? # Impatiens ! 

2 a 

■t.'amellia ! 


/ 1, Medicago ! 
/£*Ulex ! 
j(o Spartianthus. 
/ £> Pisum ! 
/b Orobus ! 
/ <& Genista ! 
/^Gytisus ! 
Lotus ! 
/ f *Rosa ! 
/7 *Kerria ! 
u *Spira3a ! 

Fragaria ! 
>. *Crataegus ! 

( lydonia. 
•i *Pyrus ! 

Eriobotrya ! 
>i *Amygdaliiri ! 
fi *Prunus ! 
J. fc*Myrtus ! 
-*7 *Punica ! — J^co*> 
/^Philadelphua ! 
/ g*Deutzia ! 

*Fuelisia!) „ ,/j 

letialf 1$ (Ik*>r*+fL*-4^LA mm . 
Clarkia ' 

/3 )kL£,i^A^vi>C<fiJL 



JL4&U if Portulaca! 

YkyvCGL * 

Gardenia ! 
Lonicera ! -^ 
Sambucus. -^ * 
Viburnum. 2- 
*Campanula ! v • 
Platycodon ! * i 
Calluna ! J^t-e^j 
Azalea ! 
Rhododendron ! 

* Arbutus ! 
*Erica ! 

*Anagallis ! J *- 
*Primula ! 3 "2- 
*Jasininum ! 3*3 

Syringa! 33 
*Vinca ! 
*Nerium ! 

Allamanda ! 

Tabernsemontana. 'i 
*Calystegia ! 

Convolvulus! ... 

Ipomsea. <> 
l*uQ« *Datura ! J fe, 

*Petunia ! - 

Solanum ! S^ 

Orobanche. 3 "J 

Gentiana. ^. 

Mimulus. m 

* Antirrhinum ! 
Gratiola ! , , 

*Digitalis ! ? * 

*Linaria ! 
Veronica ! 
Calceolaria ! ' 
Achimenes. 3 ^ 
Gloxinia ! / / 
Clerodendron l^C 
Bignonia. •&/ 
Cyclamen ! ^-2— 
Mirabilis. <0 J2L 
Lauras ! 
Gladiolus ! 
Crocus ! 
Iris ! 

*Galanthus! ^6 
Leucojum ! 
Sternbergia ! • 
Hippeastrum. <■ 

*Narcissus ! »/ 

*Orchis ! 9& 
Catasetum ! n 
Hydrocharis. <=//^ 
Asphodelus. <*/ q 

# Tulipa ! 

*Convallaria ! ' 
Fritillaria ! 

*Lilium ! 

*Hyacinthus ! ' * — 

*Polianthes ! 

*Hemerocallis ! ¥- a 

*Colchicum ! ' t 

*Sagittaria ! 

*Tradescantia \^0 
Commelyna ! > / 

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 ap- 
pearance to the combination of the following appear- 
ances — a petal-like form of the stamens, increase in 


the number of these organs and similar changes affect- 
ing the pistils, and is applied to several distinct condi- 
tions. 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 deve- 

In double flowers of Primmla 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 ! Alcea. 

*Anemone ! Hibiscus ! 

Nigella. Amygdalus ! 

*Papaver ! Loriicera ! 

*Dianthus ! Scabiosa. 

Saponaria ! iEschynanthus ! 

Viola ! Primula ! 

Camellia ! 

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 fre- 
quency among Cruciferce. 1 

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 

1 See Baillon, ' Adansonia,' iii, p. 351, tab. 12, Sinapis. 


point of view. C. Morren lias given the name adeno- 
petaly to a case wherein one of the glands at the base 
of the petals in Lopezia was replaced by a petal. 1 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. 9 

Staminody of the sepals and petals. — In the first named this 
is of very rare occurrence. M. Gris has recorded an 
instance in Pliiladelplms speciosus* which appears to be 
the only case on record. The corresponding change in 
the case of the petals is far more common. De Can- 
dolle cites in illustration of this occurrence flowers of 
the common haricot, in which the alee and carina of the 
corolla were thus changed/ 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 Gapsella bursa- 

Cramer figures and describes a stamen occupying 
the place of a petal in Daucus Garota. 5 Turpin 6 de- 
scribes a similar occurrence in Monarda fistuhsa, 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, 
Aspjliodelus ramosus, as having presented this change, 
and Wiegmann 7 has seen anthers developed on the 
awns of Avena chinensis. In semi-double flowers of 
Ophrys aranifera and Orchis maseula, the lateral petals 

1 ' Bull. Acad. Belg.,' xvii, part i, p. 516, c. tab., and 'Lobelia,' p. 83. 

2 Cited in ' Bull. Soc. Bot. France,' xiv, p. 253 (' Rev. Bibl.'). 

3 ' Bull. Soc. Bot. Fr.,' 1858, p. 331. 
* ' Mem. Legum.,' p. 44. 

5 ' Bildungsabweicb, ' Pflanz. Fam..' tab. 8, f. 12. 

ri -Atlas etc Gothe,' p. 55, t. I, f. 18. 

: Wiegmann, ' Bot. Zeit.,' 1831, p. 5, tab. i. 

B1 LMIN0D1 . 299 

are ocasionally partially antheroid, and others occur in 
which two of the outer series of stamens, which arc 
ordinarily suppressed, arc present, but in a petaloid 

slate. Keichenbach 1 figures an illustration of this 
change, and also Moggridge. 2 

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 (Alliv/m Scorodoprasv/m), 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 rusticcma), 
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* and in Genti- 
linn camvpestris.* 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. 5 

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 
mMathiola mcana in which the carpels were disunited, 
and antheriferous at the margin. 6 

The passage of pistils to stamens in willows has been 
frequently remarked, as in SalLv babylonica, sHesiaca t 
ciiierea, Gwprea 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 

1 ' Ic. Flor. Germ.,' xiii, tab. 112, cccclxiv. f. 2. 

- Seemann's ' Journal of Botany,' 1867, p. 317, t. 72, A (Ophrys). 

:< ' Enuni. Euphorb.,' p. 53. 

4 ' Limiffia,' i, p. 457. 

' - De Balsam,' p. 17. 

6 B. Clarke, 'Arrangement of Phaenog. Plants.' p. J- 


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 

Campanula persicifolia, C. rajmnculoides, and 0. 
glomerata have been observed to present an anther 
surmounting the pistil. 1 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 malforma- 
tion in Cliamcero'ps humiUs, 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. 2 

In Tqfieldia cahjculata a similar substitution of a 
stamen for a carpel has been observed by Klotsch, 3 
and "Weber 4 gives other instances in Primus and Pceoma. 
Corresponding alterations may be met with in culti- 
vated 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 
Eicinus communis as having been in one instance 

1 See ' Engehnann,' p. 26, tab. 3, f. 10, 11, 14. 

2 ' Ann. Sc. Nat.,' ser. 2, t. viii, 1837, p. 58. 

3 ' Bot. Zeit.,' 4, 1846, 889. 

' Verhandl. Nat. Hist. Ver. Preuss. Rheinl. und Westph.,' 1858, 
1860, p. 381. Cramer also, ' Bildungsabweich,' p. 90, cites a case in 
Pceonia where the carpel was open and petaloid, and bore an anther on 
one margin, and four ovules on the other. 


anthcriferous. 1 Moggridge figures a flower of Opl 
msectifera in which the rostellate process was replaa <l 
by an anther."' 

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. 

Stamuiody 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, 
X riwm Oleander, the rays of Passiflora, the corona of 
Narcissus, have all been observed occasionally to bear 
anthers. 3 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. mcompara- 
bili.s, and X. 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, 4 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 them- 
selves accidentally the characters proper to stamens. 

1 'Eupliorbiaceae,' p. 205. 

3 Seemanus ' Journ. Bot.,' iv. p. 168, tab. 47, f. 1. 

3 Moquin-Tandon, 1. c, 220, Passiflora. Masters, 'Joum. Linn. Soc.,' 
1857, p. 159, Saponaria. Seeinann's ' Joum. Botany,' vol. iii, p. 107, 

4 ' Bull. Soc. Bot. Pr.,' 1857, p. 452. 



Pistillody of the perianth. — The passage of the segments 
of the perianth into carpels has been observed fre- 
quently in Tulvpa Gesneriana, the change in question 
being generally attended by a partial virescence. M. 
Gay is said by Moquin to have observed a flower of 

Fig. 160. — Flower of tulip, showing 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. 

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. 

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. 1 

1 ' Gardeners' Chronicle,' 1866, p. 897. 



Pistillody of the stamens. — This change whereby the sta- 
mens 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 
- 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 
jrigida 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 supe- 
rior, 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. 

In some varieties of the orange, 
called by the French " bigarades 
cornues," the thalamus of the flower, 
which is usually short, and termi- 
nated by a glandular ring-like disc, is prolonged into 
a little stalk or gynophore, bearing a ring of super- 
numerary carpels. These carpels are isolated one 

Fig. 161.— Super- 
numerary carpels in 
the orange, arising 
from substitution of 
pistils for stamens. 



from another, and are formed by the transformation 
of the filaments of the stamens. 1 

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 transforma- 
tion 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.) 3 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. 3 Dunal and Campdera have 
described flowers of Rumex crisjms, with seven pistils, 
occupying the place of as many stamens. 

In Palaver bracteatum a considerable number of the 

Fig. 162. — Substitution of carpels for stamens in Papavcr. 

stamens sometimes become developed into pistils, 

i Maout, ' Lecons Element.,' vol. ii, p. 488. 

2 Poiteau and Turpin, ' Arb. Fruit,' t. 37, and Trecul, ' Bull. Soc. Bot, 
France,' vol. i, p. 307. 

3 Clos, ' Mem. Acad. Toulouse, ' 5 ser., vol. iii. 


especially those which are nearest to the centre of tlio 
Hewer, and in these flowers tlie filaments are said to be- 
come the ovaries, while the anthers are curled go as to 
resemble stigmas. A similar change is not infrequent 
in Pajpaver somniferum. Goeppert, who found numerous 
instances of the kind in a field near Breslau, sa}'S the 
peculiarity was reproduced by seed for two years in 
succession. 1 Wigand (' Flora,' 1856, p. 717) has noticed 
among other changes the pistil of Genticma Amarella 
bearing two sessile anthers. Polemoniv/m ccerulevm is 
another plant very subject to this change. Brongniart 2 
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* Bv artificial fertilization aL Brono-niart ob- 
tained fertile seeds from the central normal ovary as 
well as from the surrounding metamorphosed stamens. 
Gh&iranthus Ckeiri has lonp; 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 Chevranthus : — "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 may be soldered 
too-ether, 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 placenta?, corresponding to 
the lines of junction of the staminal carpels. 

1 ' Bot. Zeit.,' 1850, t. viii, pp. 514, 664. 'Flora.' (B. Z.) 1S32, t. xv, 
p. 252; also cited in 'Ann. des Serves et des jardins, 5 vi, pp. 241-5. See 
also Schleehtendal. ' Bot. Zeit..' 1845, t. 3, p. 6. 

- ■ Bull. Soe. Bot. France," t. viii. p. 453. 




In the accompanying figures (fig. 163, a — d) the nature 
of this change is illustrated. In some of the speci- 
mens it is easy to see that the two shorter stamens 

Fig. 163. — Cheiranthus Cheiri, var. gynanthems. a. Sepals and petals 
removed to snow carpellodic stamens, b. The same laid open. c. 
Transverse section, d. Plan of flower with four carpel-like stamens, &c. 

undergo the change into carpels later and less per- 
fectly 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. 1 

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 

See also Allmann. "Rep. Brit, Assoc.,'' July, L85] 



Fig. 164.— Struc- 
ture lialf anther, 
half carpel, IAlium. 

slmws one of the intermediate organs from these 
Bowers, in wliicli half the structure seems devoted to 
the formation of ovules, while the other half bears a 
one-celled anther. Lindley 1 has also 
described a case of this kind in a 
species of Amaryllis. 

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 in- 
stance there is usually no interme- 
diate condition between the stamen 
and the pistil. Guillemin 2 also de- 
scribes 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. 3 

In Bosa arvensis similar transformations have been 
observed of a slightly more complex character than 
those just mentioned, and passing into more impor- 
tant changes, especially to the formation of pollen 
within ovules, formed on the edges of an open car- 
pellodic 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 cer- 
tain fleshy appendages surrounding the androecium 
were provided with ovules/ 

1 ' Theory of Hoi-ticnlture,' ed. 2, p. 82. 
; ' Mem. Soc. Hist. Nat. Paris,' i, 16. 

3 Gen. Muuro, ' Trans. Linn. Soc.,' xxvii. p. 7. 

4 ' Bull. Soc. Bot. Fr.,' 1857, p. 21. 


Payer, in his ' Organogenies p. 38, mentions a 
stamen of Dioncra bearing not only an anther, but 
likewise an ovule. 

Sern/pervivwm tectorwm and 8. monfanum have long 
been noticed as being very prone to present this change. 
Mohl 1 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 

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 mal- 
formed 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 

1 _' 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. 


into a long style, terminated by a flattened Bpathulate 

phirwivm elatwm is one of the plants in which 
this change has been mosl frequently noticed. 1 

In willow- the change of pistils into staminal organs 
has been frequently observed. In SclUa babylonica 

Prof. Schnizlein has described various transition st;i^ 

Pig. 165. — . v ". tecotorwn. a. Normal stamen, h. Xorinal 

carpel. 6, c, e.f. <j. Structure partly staminal, partly carpellary. <}. 

Transverse section through c, showing pollen internally, ovules ex- 

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 Popidus.* Mr. Lowe also records the conversion 
of stamens into ovaries in 8aUx Andersoniana, and 
this by every conceivable intermediate gradation. 3 

1 Godron, ' Bull. Soc. Bot. Fr..' xiii. p. 82, Rev. Bibl. 

: Cited in Heniivy. ' But. Gazette,' iii. p. 12. 

■ • Ann. Xat Hist.." September. 1856, p. 56. See also Kirschleger, 



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 theSempervivum, 
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 spe- 
cialised structure of plants renders such a change in 
them far more easy of comprehension. 

Thalictruni minus. 

Delpbiniuin elatum. 

Magnolia fuscata. 

Bocconia cordata. 
*Papaver bracteatuin ! 
somniferuni ! 

Dionsea muscipula ! 

Barbarea vulgaris. 
*Cheirantbus Cheiri ! 

Cochlearia Armoracia. 

Tropseoluru majus. 

Citrus Aurantium. 
*Serupervivura tectoruni ! 

Begonia frigida ! 

Cucumis, sp. 

Cucurbita Pepo. 

Pyrus Mains. 

Rosa arvensis ! 

Saxifracja crassifolia ! 

Myrtus, sp. 

Campanula rapunculoides. 

Polemonium caeruleum. 

Gentiana Amarella. 
*Erica Tetralix. 

Stacbys germanica. 

Primula acaulis. 

Rnmex crispus. 
*Salix, sp. plur. ! 

Euphorbia esula. 


Aspbodelus ramosus. 


Lilium tigrinum ! 

*Tulipa Gesneriana ! 

var. cult, plurim. ! 


Zea Mays. 

Bambusa, sp. 

PistillocTy of the ovule. — An instance of this extraordi- 
nary transformation in the carnation, as observed by 
the Rev. Mr. Berkeley, is given at p. 268. 

' Flora (Bot. Zcit.),' xxiv, 1841, p. 340, Salix alba. Henscbel, * Flora 
(Bot. Zeit.),' 1832, t. xv, p. 253, 8. cinerea. Hartmann, ' Flora (Bot. Zeit.)/ 
xxiv, p. 199, S. nigricans. Meyer, C. A., ' Bull. Pbys. Matb.,' t. x, 8. alba. 



'I'm hue 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 par- 
tial perversion in organization, the former differ from 
the normal standard, not merely in degree, but abso- 
lutely. 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 hy- 
pertrophy 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. 



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, Hyper- 


trophy, Atrophy, &c, for malformations capable of 
more rigid classification than those here alluded to. 

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 

Fig. 166. — Portion of the under surface of a cabbage-leaf, with horn- 
Uke excrescences projecting from it. 

some portions as contrasted with others, whence arises 
either a depressed cavity, as in the case of a leaf, or 

Ax 1 1 HA. 


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 Nelwm- 
hiuni, 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. 

Fig. 167. — Lettuce leaf, bearing on the back a stalked cup, arising 
from the dilatation of the stalk (?). 


In a variety of Codiceum variegatum a similar forma- 
tion 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-cornutum, Moore, and peraferum, Woll. 1 

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, 3 in which 
the leaves, epicalyx, sepals, and petals, were all provided 
with tubular spurs. 

In O&phalotus 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. Brauu, 
' Flora v. Bot. Zeit.,' 1835, t. xviii, p. 41, Aristolochia.) 

Tubular formations in the flower. — A similar formation of 
tubes happens in some double noAvers ; for instance, 
it is not infrequent in double flowers of Primula 
sinensis, in which tubular petal-like structures are 

1 Moore, ' Nature Printed Ferns,' 8vo edition, vol. ii, p. 154, et p. 173. 
- 'Flora (B. Z.),' 1821, vol. iv, p. 717, c. tab. 



attached to the inner surface of the corolla; sometimes 
these petaloid tubes replace.the stamens, while at other 

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. 

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. 

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 pro- 
bably to the same causes as those which induce hyper- 
trophy. Such spurs have frequently been seen on the 
corolla of Digitalis purpurea, Antirrhinum majus, 1 
Tulipa Gesneriana, and occasionally on the sepals of 

1 Chavannes, ' Mon. Antirrb.' 


Fuchsia. They are very frequent in some seasons in 
the corolla of certain calceolarias (G. floribtmda). By 

Morren this production of ad- 
ventitious spurs was called 
" Ceratomanie." 

Similar processes may some- 
times be seen in the capsules 
of Linaria vulgaris, as also in 
the fruits of some of the Sola- 
nums, quite without reference 
to the arrangement of the 
carpels, so that their produc- 
Fig. 169. — Corolla of tion seems to be purely irre- 

Calceolaria, showing irre- P'ular. 

erular tubular spurs pro- -** ' • i 

fecting from the lower hp. Morren, as previously re- 

marked, 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 

Contortion. — An irregular twisting or bending of the 
stem or branches is by no means of uncommon occur- 
rence, the inducing causes being often some restriction 
to growth in certain directions, or the undue or dispro- 
portionate growth in one direction, as contrasted with 
that in another. Hence it may arise from insect-punc- 
ture, 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 alter- 
nately to one side or another, frequently in association 



with an oblique form of the leaf. This state is some- 
1 iincs present to an extreme degree, as in sonic varieties 
of shrubs (CratcegvA, Robmia, &c.) cultivated for their 

singularly tortuous branches. 

Fig. 170. — Portion of the culm 
of a Juncus, bent irregularly. 

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 1 alludes to a case of this kind in a 
species of pine {Firms), 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 Rkeims, the trunks of the trees are con- 
torted 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 inter- 
grafted, 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 weep- 
ing ash. 

M. Fournier 2 mentions the stems of Buscus 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, origi- 
nally observed by Goethe, and figured and described by 
Jaeger; 3 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 

1 ' Bull. Soc. Bot. France,' t. vii, 1860, p. 877. 

2 Ibid., t. iv, 1857, p. 759. 

3 Jaeger, " De monstrosa folii Plicenicis dactyliferce conformatione a 
Goetlieo olim observata," ' Act. A.cad. Leop. Oar. Nat. Our.,' vol. xvii, 
suppl., p. 293, c. tab. color, iv. 


as tin- leaflets increase in size, remaining from some 
cause or other unbroken, and thus serving to restrain 
I lie growth. 

A similar irregularity of growth occurs, not unfre- 
quently, 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. 1 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 
Alstrcemeria, 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 SpiraiitJies. 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 occidentalism 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 

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, 

1 See Goethe. ' UeV»ev die spiral Tendenz." 


&c, the degree of twisting being dependent to a great 
extent on the roughness of the surface around which 
the stem twines. 1 

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 accom- 
panied by displacement of the leaves, whorled leaves 
becoming alternate, and opposite or whorled leaves 
becoming arranged on one side of the stem only. Fre- 
quently 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 fine 
round the stem, as in a specimen of Dracocephalu/m 
specwsum described and figured by C. Morren. The 
leaves of this plant are naturally rectiserial and de- 
cussate, but, in the twisted stem the leaves were curvi- 


serial, and arranged according to the — 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 neces- 


sarily — . The fraction-— hence comes regularly into 

2 2 3 5 

the ■=■ series (j , -, — V 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 Dvpsams fullormm. (See 'Proceedings of 

1 See Darwin " On Climbing Plants," ' Jonrn. Linn. Soc. Botany,' 
vol. ix, p. 5. 



the Linnean Society, 5 March 6, 1855, vol. ii, p. 35 
The stem was distended, and hollow, and twisted od 
itself; its fibres, moreover, were ar- 
ranged in an oblique or spiral direc- 
tion; the branches or Leaf-stalks, which 
usually are arranged in an opposite and 
decussate manner, were, in this c 
disposed in a linear series, <>ne over 
the other, following the line el* cur- 
vature 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 con- 
dition of the internode between the 
two leaves, then, in the teazel-stem 
just described, each turn of the spiral 
would represent a lengthened inter- 
node ; and, if the fibres of this speci- 
men 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 expla- 
nation is borne out by the similar 
twisting which takes place so fre 

Fig. 172.— Twist- 

1 ", ed stem of Dipsacus 

quently m the species or (xalvu/m and fulhnum. 
other RuMaeecB. 

Gr. Franc 1 was one of the first to notice this twisting 
in Galium, and M. Duchartre, 2 in mentioning a similar 
instance, gives the following explanation of the appear- 
ance which will be found to apply to most of these cases. 
In the normal stem of Galium Mollugo the branches 

1 'Epbem. Nat. Cur..' dee. 2, ami. 1. 1083. p. 68, fig. 14. 

2 'Ann. des Sr-ienc. Nat.,' third Beries, vol. i. 1S44. p. 2HJ. 



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 con- 
firms 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 culti- 
vated 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 

Fig. 173. — Stem of Galium spirally twisted. From a specimen com- 
municated by Mr. Darwin. 

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 JEJqui- 
setum, 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 Eeinsch 1 cites 
a case in Eqidsetum 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 

1 ' Flora,' Feb. 4, 1858, p. 69, tab. ii, f. 3, and also ' Flora.' 1860, p. 737, 
tab. vii, f. 9. 



the two preceding, it was seen that the spire included 
the constituents of seven ordinary verticils. 

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- 

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." 1 

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 in- 
herited in the seedling plants. 

The following is a list of the plants in which spiral 
Torsion of the stem or branches has been most fre- 
quently observed. (See also under Fasciation and 

Hesperis matronalis. 
Dianthus barbatus. 
Pyrus Malus. 

Cercis siliquastruni ! 
Punica Granatum. 
Robinia pseudacacia ! 
Rubia tinctorum. 
Dipsacus fullonum ! 
Scabiosa arvensis. 
*Valeriana officinalis ! 
dioica ! 
Galium aparine ! 
* Mollugo ! 

verum ! 
Hippuris vulgaris ! 
Veronica spicata. 

Hyssopus officinalis. 
Thymus Serpyllum. 
Lamium purpureuni ! 
Dracocephalum speciosuni. 
Mentha aquatica. 

Mentha viridis. 
Fraxinus vulgaris ! 
Sauibucus nigra. 

Rumex, sp. 
Ulmus campestris. 
Casuarina rigida. 
Abies excelsa ! 
Li Hum Martagon ! 
* Asparagus officinalis ! 
Sagittaria sagittifolia. 
Epipactis palustris. 
Triticum repens ! 
Lolium perenne ! 
Phleum pratense. 
Juncus conglomeratus ! 
Scirpus lacustris. 
Equisetum Telmateia. 



arvense ! 

Among the more important papers relating to this 
subject may be mentioned : 

Moquin-Tandon, ' El. Ter. Yeg.,' p. 181. Kros, ' De Spira in plantis 
conspicua.' Morren, ' Bull. Acad. Roy. Belg.,' 1851, torn, xviii, part i, 
p. 27. Milde, ' Nov. Act, Acad. Leop. Carol. Nat. Cur., 1839. Ibid., 
vol. xxvi, part ii, p. 429, Equisetum. Irmisch, ' Flora,' 1S58, t. ii, 
Equisetum. Yrolik, ' Nouv. Mem. Instit. Amsterdam,' Lilium. Schlech- 
tendal, ' Bot. Zeit.,' xiv, p. 69, et v, p. 66. De Candolle, ' Organ. 

1 ' Bull. Acad. Belg.,' t. xvii, p. 196, " Lobelia," p. 53, c. tab. 


Veget., t. i, p. 155, tab. xxxvi, Mentha, &c. Alpli. de Candolle, ' Neue 
Denkschr. Allg. Scliweiz. Gesellschft.,' band v, tab. vi, Valeriana. Du- 
cbartre, '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 oc- 
currence as the corresponding condition in the stem. 
In Alstrcemeria 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 Oodiceum 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. Scolopendrwm vulgare var. spirale. 1 

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 occasion- 
ally springing from the sepals or petals, as indeed 
happens normally in Hodgsortia, Strophanthus, &c. 

M. Decaisne 2 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 Cobcea 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 
scabet. 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 Modecea and 
some PassifloracecB. 

1 Moore, ' Nature-printed Ferns,' 8vo edition, vol. ii, p. 183. 

2 '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. 



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 ex- 
tremities, 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 

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 terato- 
logical occurrence, though it is not easy to account 
for it. 

Fig. 175 shows an instance of the kind in a radish, 


Fig. 175. — Interrupted growth 
of Radish (from the ' American 

Fig. 176. 
iu Apple. 

-Interrupted growth 

. 176 a similar deformity in the. case of an 
the dilatation of the flower-stalk below the 

and fi 


ordinary fruit producing an apJDearance as if there were 

two fruits one above another. 

In leaves this peculiar irregularity of development 
is more common. 

328 HETER0M011PHY. 

In some varieties of Codiceum variegatuvi 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 varia- 
tions may be found in ferns, especially Scolopendrium 
vulgar e. 

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 |xj , 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. Muller, 
p. 255. 

Gornute 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 Codiceum 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, Coceoloba, Bauhinia, &c. The same 
thing occurs in the leaf-like branches of Ruscus, the 
flower- stalks of Xylophylla, Phi/Hart thus, Ptcrisanthes. 
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 

Mr. Bennie 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. 1 

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 develop- 
ment and in the position of rest. 2 

In some water plants, as Sagittaria, Alisma, Pota- 
mogeton, &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 0. linearifolium, 
in which, owing to the suppression of the lamina, the 
petiole had become dilated into a ribbon-like expan- 
sion — deformation rubanee of Moquin. 



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 

1 Loudon's ' Magazine Nat. Hist.,' vol. ii, p. 463. 

2 0. Morren, ' Bull. Acad. Belg.,' 1852, t. xix, part iii, p. 444. 


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, some- 
times 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 consider- 
able difference in the form of the leaf-organs, accord- 
ing to the place they occupy and the functions they 
have to fulfil ; but, in addition to this, it not unfre- 
quently happens that the leaf-organs in the same por- 
tion 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 tho yellow jasmine, almost 
every intermediate stage may be traced from an ovate 
entire leaf to one very deeply and irregularly stalked. 
BroussoneUia papyrifera, and Lav/rus Sassafras, and the 
species of Panax, may be mentioned as presenting 

Fig. 177. — Syringa persica laciniata, showing polymorphous leaves. 

this condition. Sometimes in the last-named genus, 
as also in Pt&ridophyllwm, every gradation between 
simple and compound leaves may be traced. The 
horse-radish (Gochlearia Armoracia) may also be in- 
stanced as a common illustration of polymorphism in 


the leaves. In ferns it is likewise of frequent occur- 
rence, markedly so in Scolopend/H/u/m 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. 

A similar protean state, but little less remarkable, 
occurs in many of our British ferns, notably in Scolo- 
pendrw/m vulgare, of which Mr. Moore enumerates no 
fewer than 155 varieties, 1 many of the forms occurring 
on the same plant at the same time. Cultivators have 
availed themselves of this tendency to produce multi- 
form 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 in- 
stance, 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 stipulated, 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, 

1 ' Nature-printed Ferns,' 8vo edition, vol. ii, p. 197. 



Pteris qtLctdriav/rita 9 in which the fronds emerging from 
an adventitious bud are very differenl from the ordinary 


Fig. 178. — Portion of a frond of Pteris quadriaurita, with an adventi- 
tious 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 con- 
stituent 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 1 {Solarwm tuhcrosvm), which produces first double 
and sterile flowers, and subsequently single fertile ones ; 
the other portions of the plant do not differ much. 

1 'Gard. Ohron.,' 184:,, p . 790. 


Stackhousia juncea, according to Clarke, has mixed 
with its perfect flowers a number of apetalous blossoms 
destitute of anthers. 1 

This peculiarity is well exemplified in the tribe 
Gaudichaudiece of the order Maljpighiacece. 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 car- 
pels 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. Violacece, GamparmlacecB, &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 com- 
pletely 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 

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, with- 
out 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 

1 ' A New Arrangement of Phamog. Plants,' p. 36. 

2 'Bull. Soc. Bot. France.' 185fi. t. iii, p. 5fi!». 


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 lias demonstrated that the di- and 
trimorphic forms of Prvm/ida, of I/wmm, Lyth/rum, and 
other plants — forms differing mainly in the relative 
length of the stamens and styles, are also connected 
with st liking 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. 1 

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 varia- 
tions end and malformations begin. There are, how- 
ever, two or three cases cited by Mr. Darwin 2 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 

1 The reader will find an abstract of Mr. Darwin's views in bis work 
on tbe ' Variation of Animals and Plants,' vol. ii, p. 181. 
5 Loc. cit., i, 336. 


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 

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 expla- 
nations 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 refer- 
ence 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 trans- 
mitted, 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 Ani- 
mals and Plants,' i, p. 373, where numerous references 
are given, and wherein certain well-known and highly 
remarkable instances, such as the Cytisus Ada/mi, the 
trifacial orange, &c, are discussed. 

UjTERATTONS of dolour. 


D I 


Changes in the colour of tin- 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 un- 
usual distribution of the solid or fluid matters on which 
the colour depends. The superposition of cells con- 
taining 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 forma- 
tion 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 
Bichardia cethiopica one or more leaves become occa- 
sionally as white as the spathe is usually. 

1 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 



Virescence. — Engelmann 1 pointed out that, so far as 
flowers were concerned, there are two ways in which 
they assume a green colour, either by a simple develop- 
ment 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 — frondes- 
cence. Morren 2 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 dis- 
tinguish 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 in- 
creased when they are subject to this change ; instead 
of falling off speedily they become persistent when so 

Some flowers are more liable to virescence than others. 
The common honeysuckle, Lonicera Perichjmenv ,,/, 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 frondes- 
cence ; for, in this instance, we have the most folia- 
ceous 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 

' ' De Antholys,' p. 32. § 38. 

2 "Bui]. Acad. Bdg..' xvii, part 2, p. 131, <■. tab. 


abortive anthers, and the pistil also is in a rudimentary 
condition. UmbellifercB are not unfrequently subject to 
this change, e. g. 9 Torilis Anth/risws, Dun, -us Carota t Hera- 
ch wm 8phondyli/u/m,Ga/ru/m carui, &c. Prvnmlacece, again, 
are frequently subject to virescence. Among Gompositoe 
the following species are recorded as having had green 
flowers — Oirsi/wm tricephalodes, Senecio vulgaris, Galen- 
dnda officinalis, Pyrethrurn Parthenium, Garckms crispus, 
Hypochceris radicata, Hieracium prealtum, Cirsium 
a/rvense s ( 'oreopsis Drummondi. 1 In Kanunculacece vires- 
cence has been observed in Delphmwm elahi/m, crassi- 
caule and Ajacis, Anemone hortensis and nemorosa, Aqui- 
legia 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 corona/rid and hortensis 
not unfrequently assume the coloration usually con- 
fined to the parts of the perianth. A similar illus- 
tration 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 

1 See Cramer, ' Bildungsabweich,' pp. 17, 55, 82, 65. See also Lucas, 
' Verliandl. des Bot. Vereins. Brandeub.,' heft 1. 2. AncJmsa. Christ. 
' Flora.' 1867. pp. 376. tab. 5, 6, Stachys. 


the petals, the others being of their ordinary green 

The occurrence of coloured bracts, as in Pomsettia, 
Bougaw/villea, &c, is very common under natural con- 
ditions, and need not here be further alluded to. 

Increased intensity of colour often accompanies 
1 1 ratological changes ; an instance has just been alluded 
to in the Gesnera : the feather hyacinth, Muscari 
cnmosum, furnishes another illustration, the adven- 
titious 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.' 



To a certain extent the number of the organs of a 
plant is of even greater consequence for purposes oi* 
classification than either their form or their arranere- 
inent ; 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 mono- 
cotyledons 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 

Considered teratologically, the changes, as regards 
the number of organs, ai B readily grouped into thos< 

342 NUMBEll. 

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 maybe, as hap- 
pens in what is termed prolification. This formation of 
buds occurring, as it does, often in unwonted situations 
is treated of Tinder the head of alterations of arrange- 
ment, the mere increase in number being considered 
of subordinate importance as contrasted with the 
altered disposition (see p. 100), 



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 trans- 
mutation 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 lacinia- 
tion 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 augmen- 
tation 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 theo- 
retical explanation, is shown by some of the Primulacece, 
wherein a second row of stamens is occasionally present 
in the adult condition, and renders the floral symmetry 

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 Delphi itium, as shown by Braun, 1 the stamens and 
carpels are members of a continuous spiral series, and 
in the double balsam an extra corolline whorl is pro- 
duced, 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 
refernble to an absolute increase in the number of 
whorls, and not to chorisis. 2 

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 unfor- 
tunate 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, 

1 Braun, ' Pringsheim Jahrbuch f. Wiss. Bot.,' 1858, 1, p. 307, tab. 22, 23. 
- Henfrey, 'Jour. Linn. Soc. Bot.,' vol. iii, p. 159. 

( B0EISI8. 3 I-') 

"parallel chorisis;" oral right angles to it ••collateral 
chorisis." Endeed, before bo much attention had been 
paid to tin- way in which the floral organs are deve- 
loped, it was thoughl that an actual splitting and 

dilamiiiation 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 pri- 
mary 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 tlie 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. 1 

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 Orchidacece, of which one 
only, under ordinary circumstances, is developed. 
When the numerical symmetry is restored, as it some- 
times is, it is obvious that the augmentation that 
occurs is of a different character from that arising from 

1 On the subject of. chorisis or dedotiblement the reader may profit- 
ably consult Moquin-Tandon, ' Ess. but les Dedoublernents,' 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. hi. p. 355, adnot. Lindloy, 
■ Elements of Botany,' p. 76. Asa Gray. ' Botanical Text Book. 9 


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, bnt if from chorisis or " dedouble- 
ment," it may happen that the normal arrangement is 

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. 



By Linne an undue number of branches was desig- 
nated 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 
Pica}, quod vulgo a genio ortum arbitrator ; frequens 
apud nos in Betula, prccsertim Norlandice, in Garpino 
Scanice, nee infrequens in Pinu." 1 

By some of the older authors this condition Avas 
called polyclady. Tn some cases, it would seem to 

1 'PhilBot..' §274. 



be duo to fungi as in the witches' brooms (hexenbesen) 
of the German forests ; in otlier instances, it is a result 
of mutilation as after the operation of pollarding. 

Mdjuin-Tandon 1 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 " burr's," 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. 

Occasionally, a great increase in the number of 
pedicels, or flower-stalks, may be met with in conjunc- 

Fig. 17!). — Flower stalks of BeUevalia comosa. nat, size, after Morivn. 

•El. Ter. Veget.,' p. 392. 



tion with a decreased number of flowers, as in the 
wig-plant (Rhus Cotiims), or the feather-hyacinth 
(Bellevalia comosa). In these cases the supernumerary 

Fig. 180. — Tuft of branches at the end of the inflorescence of Belle- 
valla comosa, enlarged after Morren. 

pedicels are often brightly coloured. To this condition 
Morren gave the name mischomany, from n'loyoq, a 
pedicel, a term which has not generally been adopted. 1 

M. Fournier 2 describes a case in the butcher's broom 
(Busgus aculeatus), wherein from the axil of the minute 
leaf subtending the flower a secondary flattened 
branch proceeded. 

Duchartre 3 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 pre- 
mature development of some of the axillary bulbils into 
flowering steins which become adherent to the primary 
flower-stalk, cannot, in all cases, be determined. Cer- 

1 ' Bull. Acad. Belg - .,' xvii. part ii. p. 38. 
: -Bull. Soc. Bot. Ft.,' vol. iv, 1857. p. 760. 
•■ Ibid., vol. viii, 1861, p. 159. 

win: ORG WS. 3 I'.i 

tainly, in some cases examined by me the hitter was 

the case. 1 

Under this head, too, may be included those eases 
wherein an ordinarily spicate inflorescence becomes 
paniculate owing to the branching of the axis and the 
format ion of an unwonted number of secondary buds. 

Fig. 181. — Increased number of male catkins in the hazel Corylus 

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 them- 
selves which is proper to the leaves. These latter, 
indeed, replaced the strobili above. 2 

M. Reichardt describes an analogous case in the 
same species, and attributes the inordinate number of 
cones to a fungus {Peri&ermivm pini). In this case 

1 See ' Gard. Chron.,' July, 1866, p. 656, and Clusius, ' Plant. Rar.,' 
lib. 2, p. 143, Tulipa scroti na irv\vK\aci)q, minor, &e. ' Hort. Eysttett. 
Plant. Vern.,' fol. 12. 

- ' Jaeger de Pini sylvestris monstrositate,' Stuttgardt, 1828. 


there were no less than 227 cones, but each one half 
the size of the ordinary cones. 1 

Of a similar character is the many -headed pine-apple. 
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 (Triticwm comjiositum) , 
certain varieties of Maize, etc. Similar exuberant 
growths occur in OrcMdacece, in Oyperacece, e. g. Carex, 
in EestiacecB, 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 Equisetacece a similar plurality of spikes 
occurs often as a result of mutilation. 2 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 
cestwum described by M. Gay. 3 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 

i Cited in ' Bull. Soc. Bot. Fr.,' xiv, p. 265. 

2 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. 

3 'Bull. Soc. Bot. Fr.,' vi, 266, vii. 457. Irmisch. f Knollen und 
Zwiebelgew.,' tab. 7, figs. 10, 11. 


From the instances cited it is clear that branching 
of the inflorescence occurs most frequently in those 
plants naturally characterised by a dense compact 

Fig. 182. — Broccoli, with six perfect heads on one stalk (' Gard. 
Chron.,' 1856, Oct, 25). 

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). 

Increased number of florets in the individual spikelets of 
grasses is also met with under some circumstances. I 
have seen this in Hordcum and Lolium, and an instance 
is figured in Arena by Dr. Wiegmann. 1 M. Duval Jouve 2 
records a similar occurrence in Catabrosa aquatica, the 
spikelets of which contained from two to seven flowers. 3 

1 'Flora,' 1831, p. 5, tab. i; see also Hanstein, 'Flora,' 1857, p. 513. 
Schlechtendal, ' Bot. Zeit.,' xviii, p. 381. 

2 ' Bull. Soc. Bot. Fr.,' ix, p. 8. 

3 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 forma- 
tion (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 




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 juxta- 
position, yet originate from different parts of the stem, 
but by coalescence or lack of separation produce the im- 
pression 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 distribu- 
tion 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 

of the sub-division of the leaf-like organs of Sciadopitys vcrticillata. In 
one instance the pseudo leaf divided, and from the division proceeded a 
little axis, bearing at its summit a verticil of psendo leaves. This divi- 
sion 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 Horti- 
cole,' 1867, and 'Report. Bot. Congress.' London, 1866, p. 124. 



>in face of which project, at right angles to the primary 
plane, other secondary leafy plates; but these are, 
strictly speaking, cases of hypertrophy (see Hyper- 

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 " polvphylly," 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. 

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 

Fig. 183. — Supernumerary leaflet, Uhnus campeslris. 

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 

Fig. 184. — Supernumerary leaf, Ulmus montana. 

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 

I'm, I \i; ORG w-. 


tubular, born-like projections, and in the centre a 
thread-like process representing the midrib and termi- 

Fig. 185. — Supernumerary leaf of hazel. 

nated by a small two-lipped limb. 1 Dr. Ferdinand 
Muller speaks of a leaf of Pomaderris elliptica as bearing 
a secondary leaf on its under surface. 3 

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 "autopkyllogenv.'" 
The Belgian botanist figures a small perfect leaf spring- 
ing from the nerves of the upper surface of the primary 
leaf in a species of Miconia. As in the hazel, the direc- 
tion 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 

1 'Bull. Soc. Bot. Pr., s vol. vii, 1860, p. 587. 

2 ' Fragment. Phyt. Austral.,' part xx, p. 270. 

'Bull. Acacl. Belg., s xvi. pt. i, p. 60, "Fuchsia," p. 125, c. ic 


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. 1 

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 adven* 
titious leaflets are found almost as frequently as four. 

Walpers describes a case where the leaf of T. r&pms 
bore seven leaflets. Schlechtendal alludes to a similar 
increase in number in Gyiisus 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. TaberuBemontanus 
■ Krauterbuch,' S. 222. Schlechtendal, ' Bot. Zeit.,' ix, p. 583, xiv, p. 71. 

i " Theorie de la feuille," ' Arch, des Sciences Bibl. Univers.,' 1868. 

POLIAB ORGANS. :> > : >7 

Maugin, ' Bull. Soc. Bot. Fr..' istiti, t. xiii, p. 279. See also Cramer, 
' Biidun^s;il>\V(.'ich.' ]). !»-. Walpers, ' Liunsea, 1 1840, p. 362 (7 -leaved). 
Schlechtendal, ' Bot. Zeit.,' 1844, p. 157, Oytiam. Wigand, ' Flora,' 1856, 
p. 706. 

Frondiferous leaves have much the appearance of 
branches provided with leaves, and bhey may be com- 
pared with those instances in which an adventitious 
bud is placed on the surface or edges of the leaves, as 
in Gesnera, Cardamme, &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. m/ultistipulata) . 

An increase in the number of the spathes has been 
often noticed in Arads. 1 Prof. Alex. Braun has studied 
this subject in some detail. 2 In Oalla pahistris 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 super- 
numerary 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 

1 See Engelmami. 'Be Aritholysi,' p. 16, section 12. 

: ' Verhamll. des Botanisch. Vereins Brandenburg, 3 1S59. 1 heft. 


occur m other Arads as Arwm maculatum, Richa/rdia 
cethiopica, and Anthwri/wm Scherzericmv/m, frequently 
combined with a leaf-like appearance of the spathes 
and sometimes with a subdivision of the spadix into 
two or three branches. 

Eno-elmann 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, Gornus mas, Vinca minor, &c. 

Paris quadrifolia may frequently be met with with 
five leaves in its whorl, or even six. 1 

Increased number of bracts. — This is not of infrequent 
occurrence ; one of the most curious instances is that 
recorded by Mr. Edwards 2 in Cerastiu/m glomeratmn, 
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 

' See Henslow, 'Mag. Nat. Hist., 1 1832, rol. v, p. 429. 

2 • Phyt< 'legist. September, 1857. 


other perceptible change, while at other times the 
number of tne other parts of the flower is proportion- 
ately increased. Inn flower of a plum Bis -'pals in 

place of five sometimes exist; a precisely similar 
occurrence in the flowers of the elder (#am&wctt«), the 

i>s>'>i, and of (Ena/nth crocata, may occasionally 
met with. In the hitter case, indeed, there are 
times as many as ten segments to the calyx, and 
this without the other parts of the flower being 
correspondingly augmented. Among monocotyledons 

similar inert not uncommon, as in Tidipa, 

Allium, Iris, . Sec. 

In some plants there seems to exist normally much 
variation in the number of parts; thus in some species 
of I in adjacent flowers the calyx may be 

found with four, five, or six segments. 

Most of the ; polyphylly affecting the calyx 

maybe explained by lateral chorisis or fission. 

Polyphylly of the corolla. — This may happen in connec- 
tion 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 suppi 
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 Balsamirn ce. In ihe common garden balsam 
the fifth petal is occasionally present, while in 1! 
triflora this petal is always present. 

In a flower of a I n recently examined there 

were ten petals in one dditional five being 

evidently due to the subdivision of the five prim 
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 
w an im cl numb- 



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 with- 
out any other visible deviation from the ordinary con- 
formation. 1 

I am indebted to Mr. Hemsley for the communication 
of a similar specimen in 0. a/pifera, 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 augmenta- 
tion 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 
(seep. 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 

Anemone ' 


Ranunculus ! 


Aconitum ! 

Cyclamen ! 


Primula ! 


Anagallis ! 

Diantbus ! 


Pelargonium ! 

Syringa ! 



Sarotliamnus ! 


Lotus ! 


Primus ! 

Convallarnt ! 


Paris ! 

(Enanthe and Uinbellif. pi. ! 
Sambucus ! 

Hyacinthu.s ! 
Allium ! 


Orcbideae, sp. pi. ! 


1 Seemann's ' Journal of Botany.' iv, p. 168. t. 47, f. 3. 


For other illustrations see multiplication of whorls, petalody ; Bee also 
Moquin, loc. tit., ]>. :>">o. Engelmann, loc. cit., p. 20, § 18. Cramer, loc. 
-it., 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, HyacmtJvus, Tulipa, Iris, 
Tigridia, &c, and more rarely in Yucca (Y. Jiexilis 1 ). 

The increased number of stamens in a single whorl 
may result from a development of organs usually sup- 
pressed, and constitute a form of regular peloria as 
in I/ma/ncbf wherein a fifth stamen is occasionally met 
with. Among normally didynamous plants such nume- 
rical restitution, so to speak, is not unusual ; thus, in 
Veronica four and five stamens occur. Fresenius has 
seen five stamens in Lamium, Mentha, Chelonef 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 3 figures a flower 
of Mattldola annua with five long stamens instead of 
four ; one of the long pairs of stamens has here under- 
gone a greater degree of repetition than usual. De 
Candolle 4 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 5 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 6 mentions the presence 

1 ' Illust. Hortic.,' 1866, misc., p. 97. 

2 See Fresenius, ' Mus. Senkenb.,' bd. 2, p. 43. Schlechtendal, ' Bot. 
Zeit.,' iv, pp. 403, 492, Veronica tetremdra. 

3 ' Flora,' 1865, tab. 6, fig. 8. 

4 ' Org. Veget.,' t. i, p. 497, pi. 42, f. 3. 

5 • El. Ter. Veg.,' p. 354. 

e Cited in "Rev. Bibl." of 'Bull. Soc. Bot. Fr./ 1866. p. 171. 


of seven stamens in another Crucifer, Ionopsidi/um 
lira ale. 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 IAquidambcvr 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 Malvacece, St'erculiacece, Buttneriacece, Tiliacece, 
and Myrtacece. It is probably the idea of splitting or 
dilamination involved in the word chorisis that has 
led many English botanists to hesitate about accept- 
ing 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 1 goes 
far to support the notion here adopted. The majority 
of the stamens of laurels (Lauras) 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 

1 Loc. cit . 35] 

QYN(E< ll M. 363 

M. Clos 1 mentions a flower of rue (R/uta) wherein 

there were two stamens joined together below and 
placed in front of a petal, as in Vegam/um. 

Buchenau 2 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 Sarotha/m/rms 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 conse- 
quent 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. 

Polyphyily of the gyxicechim. — Moquin 3 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, aug- 
mentation 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 Autirrh in urn, where there are usually only two carpels 

i 'Mem. Acad. Toulous.,' vi, 1862, ex ' Bull. Soc. Bot. Fr.,' "Rev. 
Bibl.," vol. ix, 1862, p. 127. 

2 ' Flora.' 1857. p. 289. 

3 L. c, p. 354. 


present, but where, under peculiar circumstances, five 
may be found — thus rendering the symmetry complete. 1 
In Pcvpilionacece, 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, 3 in Mimosa five, in Umbelllferce three to five ; 
in some composites, e. g. Spilanthes, five carpels have 
also been noticed; in Cruciferce three and four, in 
grasses three.' 3 The double cocoa-nut affords an illus- 
tration 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 develop- 
ment 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 Cruciferce, TJmbelli- 
ferce, and Liliacece, are the orders most frequently 
affected. Cases of peloria are not included in the 
subjoined list. 



Pseonia ! 
Delphinium ! 

Citrus ! 
Prunus ! 
Amygdalus ! 
Crataegus ! 
Fuchsia ! 


Trapa ! 

Draba ! 


*Cheiranthus ! 

*Phaseolus ! 



Brassica ! 




*Acer ! 

Trifolium ! 

1 Giraud, ' Ed. Phil. Mag.,' Dec, 1839. 

2 See Cerasus Caproniana, D. C. ' Plant. Rar. Hort. Genev.,' tab. 18. 

3 Nees, ' Linna3a,' v, p. 679, tab. 11 (Schcenoclorus). 





Passiflora ! 
D auciis ! 
Angelica ! 
Heracleum ! 
Campanula ! 
Chrysanthemum . 
Primula ! 
Fraxinus ! 
Datura ! 
Solanum ! 

Veronica ! 
*Digitalis ! 
Antirrhinum ! 
Gloxinia ! 
Mercuriali s ! 
Corylus ! 
Cocos ! 
Tulipa ! 
Iris ! 

Narcissus ! 
Allium ! 
Gagea ! 
Tradescantia ! 

A few additional references may here be given to papers where an 
increased number of carpels is described : — Engelniann, ' De Antholys.' 
§ 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, 
pluricarjoellary Crucifers. Weber, ' Yerhandl. Nat. Hist. Yereins. 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 legumi- 
nous plants with polycarpellary pistils. Munro, Gen., ' Linn. Trans.,' 
vol. xxvi, p. 26, Bambusea*. Alph. de Candolle, ' Neue Denkschrift,' 
Cheiranthus. Schimper, ' Flora,' 1829, ii, p. 433. Wigand, ' Bot. TJnter- 
such.' Fleischer, ' Missbild. Cultur Pfl.' Cramer, ' Bildungsabweich,' 
p. 65, TJinbeUiferw. 

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 1 in a plum, wherein there was an 
increased number of sepals, a corresponding augmen- 
tation 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. 

Fig. 186. — Plum. Increased number of parts in the calycine, corol- 
line, and carpellary whorls respectively. 

In Pr'tmulacea? this general augmentation has been 
frequently noticed. 2 

Among Orchideoe the instance related by Dr. Seu- 
bert is worth alluding to here. This botanist observed 
and figured a flower of Orchis palustHs with tetra- 
merous arrangement of parts, that is to say there were 

1 ' Gard. Ohron.,' 1852.. p. 452. 

- See Cramer, ' Bildungsabweich,' pp. 16, 21. 

0V1 I.! s. 

four outer segments to tin* perianth, four petals, of 
which two wore lip-like, four stamens, three of which 
were rudimentary, and an ovary with four parietal 
placentas. 1 

The following list will serve to show in what plants 
this general augmentation of parts has been observed 
most frequently : 

Ranunculus. Sambu i 

Clematis ! *Priniula ! 
Delphinium. Anagallis ! 

Brassica ! Lyciuru. 

Ruta. Solanum. 

Acer ! Symphytum. 

Prunus ! Syringa ! 

Rosa ! Linaria. 

Rubus. Chenopodium. 

Philadelpkus ! *Paris ! 
Chrysosplenium Convallaria ! 

UmbeLUferse, sp. pi. '. Allium. 

*Puchsia ! *Lilium ! 

(Enothera. *Tnlipa ! 
Adoxa . Omithogalum . 

Bryonia. *Gagea ! 
Cucumis ! Tradescantia ! 

Campanula ! Orchidese, sp. pi. '. 

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 Prirrmla, the ovules and seeds are pro- 
duced in large quantities, it is not practicable to ascer- 
tain 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 mea- 
sure, 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 folia - 

1 • Linncea/ 1842, p. 389, c. ic. 


ceous 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 Rammcuhis 
Ficaria, 1 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 cen- 
tral 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 Tri- 
folium repens becomes foliaceous (see Frondescence), 
the outer ovules are generally two or more instead of 
being solitary. So, also, in the Rose with polhniferous 
ovules (see p. 274). Among Umbelliferce 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. 2 

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 inter- 
esting 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 occa- 
sionally 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 ordi- 

1 Seemann's ' Journal of Botany,' 1867, vol. v, p. 158. 

2 Cramer, ' Bildungsabweich,' p. 66, Astrantia major, Eryngium, to 
which may he added Dauciis, Heracleum, &c. 


nary circumstances cease to grow (sec ante, p. 364). In 
llic case of a ripe nut with two seeds it might be im- 
possible 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 

Increased number of embryos. — A ripe seed usually con- 
tains 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 Cycadece and Coniferce. In 
the seeds of the orange (Citrus), in those of some JEuphor- 
biacece, &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. 1 

Plurality of embryos lias also been observed in — 

Rapbanus sativus. *Viscum album ! 

*Citrus Aurantium ! Daucus Carota. 

Diosma, sp. Ardisia serrulata ! 

Hypericum perforatum. Cynancbum nigrum. 

Tripbasia aurantiaca. fuscatum. 

*JEsculns Hippocastanum ! Eupborbia rosea. 

Euonymus latifolius. Ccelebogyne ilicifolia. 

*Mangifera indica ! Allium fragrans. 

Eugenia Jambos. Funckia, sp. 

Amygdalus vulgaris ! Carex maritima. 

Yicia, sp. Zea Mays. 

Cassia, sp. 

1 See also Reinwardt, ' Nov. Act. Acad. Nat. Cur.,' 12, 1, 37 ; and 
Masters, ' Journ. Linn. Soc.,' vi, p. 24. 



See Schauer's translation of Moquin-Tandon, ' El. Terat. Veget.,' p. 
245, adnot., and ' Al. Braun Polyembryonie.' 

Increased number of the cotyledons. — Although the pre- 
sence 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 coty- 
ledons respectively. 

I have also seen three cotyledons present in embryo- 
plants of Correa, Crataegus Oxyacantha, Diantlms 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. 1 Jaeger 
alludes to a similar instance in Ajrium Petroselinum ; 2 
Ehrenberg to one in the marigold {Calendula) ; 3 Reinsch 
to an analogous appearance in the beech (Fagus), as- 
sociated 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. 4 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 cotyle- 
dons, as happens, according to Duchartre, 5 in some 

1 ' Organog. Veget.,' tab. 53. ■ ' Missbild.,' p. 206. 

3 Ehrenberg, ' Flora,' 1846, p. 704. * ' Flora,' 1860, tab. 7. 

5 ' Ann. Sc. Nat.,' 3 ser., t, x. p. 207. 

PLBI0TAX1 . 6i 1 

ConiferaB, which he considers to be improperly termed 
polycotyledonous. "Whether this holds good in the 
Loranths, where (Nuytsia, Psittaccmthis) an appearance 
of polycotyledony exists, is not stated. In the case of 
the rue (Ruta) figured by M. A. de Jussieu 1 this 
splitting of one cotyledon into two is sufficiently evi- 
dent, 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 Engel- 
mann, Moquin, and others, and the two seem to require 
distinctive epithets ; hence the application of the terms 
polvphylly 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 Mo3sa, and in a peculiar variety of carnation, called 
the wheat-ear carnation. 2 In some of these cases the 
increase in the number of bracts is attended by a cor- 
responding suppression in the other parts of the flower. 
Such a condition has been frequently met with in 
Gentiana AmareUa, where the bracts are increased in 

1 ' Mem. Mils.,' xii. t. 17. 

- ' Nov. Act. Acad. Nat, Cur.." xv. tab. xxviii. f. 3; ' Bot. Mag./ t. 1622. 
" Caryophyllus spicain frutnenti referens." A similar malformation in 
THanthus barhatus is not uncommon. It lias lately been introduced into 
wardens under the name of Dianthus " mausseux" but is not likely to 
find favour with crardener^. 



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. 

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 lias been noticed also in the common pea, Piswm 
sativum, and M. Lortet 1 records a case of the kind 
in Erica multijlora, the flowers of which, under ordi- 
nary circumstances, are arranged in clusters, but in 
this case the pedicels were more closely crowded than 

1 ' Bull. Soc. Bot. France,' t. vi, 1859, p. 268! 



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. 7G9, is figured 
a corresponding instance of Delphinium Consolida, in 

Fig-. 188. — Delphinium Consolida. 
Multiplication of bracts at the ex- 
pense of the other parts of the 

Fig. 189.— Multiplication of 
bracts, &c, Pelargonium, 

which the bracts were greatly increased in number, 
petaloid, and, at the same time, the central organs of 
the flower were wholly wanting. 

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 like- 
wise occasionally been observed subject to a similar 


Comas mas and 0. suecica sometimes show a triple 
involucre. 1 Irmish 2 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 3 
mentions as having produced bracts in unwonted 
numbers, Lythrum Salicaria, Plantago major, Veronica 
spicata, Ecliium vulgare, Melilotus arvensis, and Rubus 

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 exclu- 
sion 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 Reaper observed a calyx con- 
sisting of a double series, each of five sepals, in con- 
junction with other changes. 4 It is also common in 
double columbines, delphiniums, nigellas, &c. 

In the 'Revue Horticole,' 1867, p. 71, fig. 9, is de- 
scribed 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 

i "Weber, ' Yerliandl. Nat. Hist. Vereins. Rhein. Pruss.,' 1860. 
• <Bot. Zeit.,' 1848, p. 217. 
:f ' De Anthol.,' p. 17, § 12. 
4 ' Linnsea,' vol. ii, 1827, p. 85. 


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 malfor- 
mation, 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 1 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, 
Myosotls palustris, Veronica media, Aquilegia vulgaris, 
Nigella damascena, Campanula rapnnculoides. 

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 (Gonvallaria 
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 repe- 
tition or pleiotaxy, partly to the substitution of petaloid 
segments for stamens and pistils. 2 

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 

1 ' De Antholysi,' p. 17, tab. iii, f. 15, 16 ; Weinmann, ' Phytanth. 
iconogr.,' nro. 292. 

2 See Hildebrand, 'Bot. Zeit.,' 1862, p. 209, tab. viii ; Cramer, 
' Bildungsabweich.,' p. 7, tab. xiii ; Engelmann, ' De Antbolysi,' 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 


disposed in an irregular spiral manner at the extremity 
of the stem, some of the uppermost being occasionally 
verticillate. 1 

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, 2 a statement that is true in the main, though it 

1 Schlechtendal, 'Bot. Zeit.,' xx, 1862, p. 301. 

2 ' Phil. Bot,,' § 126. 


requires modification. In the case of polypetalous, or 
rather dialypetalous flowers, the petals may be vrery 
largely increased by multiplication, as in roses, ane- 
mones, 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 Legummosce, such as Ulex europmis 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. 1 

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 co- 
here one with another. Among double flowers of this 
character may be mentioned Campanula rotundifoUa, 
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." 

1 C. Morren, ' Bull. Acad. Belg./ xix, part ii, p. 17. 



The same condition occurs frequently in some 
species of Datura and Campanula. 

Fig. 191. — Campanula rotundifolia. " Double flowers" resulting from 
dialysis and multiplication of the petals. 

In Antirrliiim in mo jus 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 Injemalis. 

Mr. "W. B. Hemsley has kindly furnished me with 
flowers of a similar kind occurring in wild specimens of 

AKDBdiOIUH. 379 

Epaeris impressa* and there are analogous phenomena 
in the common honeysuckle (Lonicera Perictyirienum), 
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 con- 
fluent petalodic stamens ; that the latter is the true 
explanation, in certain cases at least, is shown by some 
flowers of Dctiwa fastuosa, in which the second corolla 
was partially staminal in its appearance, and bore 
nearly perfect anthers, in addition to the five ordinary 
Bt aniens, 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 an&rcecmm. — An increase in the number 
of whorls in the stamens is very common, especially in 
cases where the number of circles of stamens is natu- 
rally large. The augmentation of the number of sta- 
mens is still more frecnient where these organs are 
arranged, not in verticils, but in one continuous spiral 

In CruciferoB 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 Ca/ryo- 
phyUem, three and four verticils of stamens have been 
met with. In Lonicera Peridymemvm a second whorl 
of stamens more or less petalodic sometimes occurs. 

Moqnin mentions a variety of Balus frviicosus in 

1 ' Seeinann's Journal of Botany," iii. p. 354. 


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 Bitaxis 
land folia. 

Increased number of stamens in orchids, &c. — Various de- 
viations from the ordinary type of orchid structure 
have been already alluded to under the head of dis- 
placement, fusion, peloria, substitution, &c, but the 
alterations presented by the andrcecium 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 pre- 
sented by the andrcecium 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 inexplic- 
able 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 distribu- 
tion 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 

ORCIIIDAi'l 1 . 


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. 

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 Ci/pripedium, while the ordinary sta- 
men of the outer row is deficient, two of the inner 
series are present. The diagram, fig. 192, will serve 

Fig. 192. — Diagram showing the arrangement of parts in an orchid 
flower. According to Criiger, the stamens A 2, A 3, should be distinct 
from the lip. The uppermost figure 2 should have been 1. (See text.) 

to show the arrangement of the parts as above de- 
scribed, -f- 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 (in- 
correctly 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 sta- 
mens, 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 homo- 
logies of orchid flowers. 1 All that can be done in these 
pages is to set forth the evidence furnished by terato- 
logy 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 interpreta- 
tion put upon the facts thus recorded is that of the 
present writer, and not necessarily that of the original 

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 ). 2 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. 3 

Roeper is stated by Cramer 4 to have seen a specimen of Orchis morio 
with two stamens. 

In a flower of Habenaria chlorantha, described by the late Professor 
Henslow, 5 the outer three stamens are suppressed, while two of the inner 
group are present, as happens normally in Cypripedium. 

' On this point the reader will find an excellent summary in Lindley's 
' Vegetable Kingdom,' ed. iii, p. 183a, and in Darwin, ' Fertilisation of 
Orchids,' p. 292. See also Cruger, ' Journ. Linn. Soc' t. viii. p. 134. 

■ ' Seemann's Journal of Botany,' vol. iv. p. 168. tab. 47. 

3 Ibid., t. iv. 1866, p. 168. t, xlvii, f. 1. 

4 ' Bildungsabweich,' p. 8 ; see also ' Bot. Zeit..' 1852. p. 425. 
1 "Journ. Linn. Soc..' t. ii, p. 104. tab. 1. fig. b. 

OEOHIDAi ivl'. 383 

A flower of Catthya violacca 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 Dendrdbivm normale, Falconer, not only is the perianth regular, 
but the column is triandrous, 1 the three stamens (according to the 
diagram of its structure given by Lindley) pertaining to the outer row. 

In a specimen of Dendrobimn lucmoylossum 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. 2 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 0. 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." 3 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 confor- 
mation 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 4 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 

1 Lindl., " Orchid. Ind.," ' Jour. Linn. Soc.,' iii, p. 9. 

■ ' Arch. Bot.,' ii, p. 300, tab. xvi, f. 11. 

3 ' Seemann's Journal of Botany,' v, p. 318, tab. lxxii, figs. A 4. 1 

* " Monstr. Veg.," in ' Neue Denkschrift,' p. 17, tab. vii. 


conformatce) in Orchis morio. 1 Richard, as cited by Moquin-Tandon, 
Lindley, and others, desci-ibes and figures a peloria of Orchis latifolia 
with regular triandrous flowers. 2 

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 3 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 wotild 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.* 

A specimen of Catasetum ebiimeum 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. 19-1), 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 pari of the 
flower, there was the ordinary column, and at the opposite side, alter- 
nating 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 ar- 

1 * Flora,' t. viii, 1825, p. 736. 

- ' Mem. Soc. d'Hist. Nat.,' ii, 1, p. 212, tab. iii. 

3 ' Bull. Soc. Bot. Ft.,' t. vii, 1860, p. 26. 

4 ' Beitr. Morphol. und Biol. Orchid.,' quoted by Cramer ; ' Bildungs- 
abweich,' p. 9. 

()R( IIIDACE !'. 


rangeinent of the coloured spots on the time labellum, and that on tho 
adventitious lips, replacing the two lower of the outer stamens, 

Fig. 193. — Diagram showing the arrangement of parts in a mal- 
formed flower of Ophrys aranifera (see p. 384). 

not of a similar character. The supernumerary lips had the w-shaped 
marking which is so common in this species, -while the time lip was, 
as to its spots, much more like 0. 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 

Fig. 194. — Malformed flower of Ophrys aranifera with two super- 
numerary lips and three additional stamens. 

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. 1 

A similar illustration, for a knowledge of which the writer is in- 
debted to the kindness of Professor Asa Gray and Mr. Darwin, occurred 
in some specimens of Pogonia ophiocjlossoides collected by Dr. J. H. 
Paine in a bog near Utica, New York. It will be seen from the fol- 
lowing description that these flowers presented an almost precisely 
similar condition to those of the Ophrys arcmifera 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 ai*e just in 
the position of the two suppressed stamens of the outer series, viz. of 
A 2 and a 3, 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, A 1. Secondly, in one of the blossoms, and less distinctly in 
another, two lateral stamens of the inner series (a 1 and a 2) are repre- 
sented 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 

A tetrandrous flower of Cypripedium has also been recorded. 

In Isochilus, according to Cruger, there are often five stamens, and 

Fig. 11*5. — Diagram of flower of Orchis mascula with two additional 
lips, two perfect and two imperfect stamens (after Cramer). 

1 Masters, ' Journ. Linn. Soc.,' viii, p. 207. See also Rodigas, ' Bull. 
Soc. Bot. Belg.,' iv, p. 266, for similar changes in Cypripedium HooJeerce, 


there are Beveral, besides those already mentioned, in which sis more 
or Less | miens have been seen — of these the following may 

be taken as illustrations. Ahexandrous flowerof Orchis rmlitariB has 
recorded by Kirschleger, 1 and in the accompanying diagram (fig. 
195), from Cramer, 1 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. 

MoiT-en 1 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 eorolline 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 thi'ee sepals, two lateral petals partially lip- 
like in aspect, a third labelluni normal, two additional labella represent- 
ing 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 horuo- 
logue. Thus it was in a specimen of Odontoglossum Alexandra; ex- 
amined 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. Sec. 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 Orchi- 
dacece may be found. These stamens may be all perfect (polliniferous), 
or, as is more frequently the case, more or less petal-like. Moreover. 

' Kirschleger. ' Flora." 1844, p. 131. 

-' " Bildungsabweich,' p. 11. tab. xiv. f. '■). 

• ■ Bull. Acad. Roy. Belg.,' t. xix, part 2. p. 171. 



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 gene- 
rally, but not always, attended by some coincident malformation, of 
which the most frequent is cohesion of two or more sepals, and conse- 
quent 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. 

Pleiotaxy of the gynoecium. — An increase in the number 
of whorls of which the pistil consists is not of very fre- 
quent occurrence. Generally after the formation of the 
whorl ©f carpels, the energy of the growing point ceases, 
or if by chance it be continued, the result is more gene- 
rally the production of a new flower-bud (median pro- 
lification) than the repetition of the carpellary series. 
It is necessary also to distinguish between the verit- 
able 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 un- 
frequent, 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 

Fig. 196. — Increased number of 
carpels, tulip. 

Fig. 197.— Fruit of St. Valory 
apple cut lengthwise. 

QYU(E< l' 

in spira] series, is nut strictly referable to the pree 

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, bui at other times the incr< 
occurs without any alteration in the stamens or other 
tns. If the adventil bus carpels be exposed, they are 
ered with yellow rind, while those portions thai 
ered by the primary carpels are destitute of rind. 
S me varieties of the double tulip are very subject to 
a similar change, but, in this case, the petals and the 
iiens very frequently become more or less carpel- 
lary 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 

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 {Jjycop&rsicum ■ I is another 

plant in which an adventitious series is frequently 
produced, and generally in combination with the pri- 
mary ser 

In the Chinese primrose (Primula si't a super- 

numerary whorl is frequently met with, generally asso- 
ciated with other changes in the construction and 
arrangement of the parts of the flower. 

M. de Candolle 1 mentions a flower of Gentiana pur- 
with four carpels in one series, and five others 
in the circle immediately above them. Wigand 2 alludes 
to an instance wherein there was a second pair of 
carpels above the first in Vinca herba/iea. 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 flow 

1 ' Organogr. Veget.,' t. i. p. 509, tab. 40, fig.^. ,; . ~ ■ 
'- ■ Flora." 1856, p. 710. 


of the same plant for two years consecutively. In 
Aquilegia I have met with a similar increase in the 
whorls of carpels. 1 Meissner records a similar aug- 
mentation in Polygonum orientale* 

Wigand 3 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 cathar- 
tica. Eichler 4 has put on record a similar case in a 

Marchand 5 mentions a polycarpellary berberid (Epi- 
medinm Musschianum). The supernumerary carpels 
in this flower were placed on a short axis, which origi- 
nated 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 1 from a transformation, or substitution of 
carpels for stamens. 

Lastly, the instance cited by Dr. Allman 6 in 8cm- 
fraga G&wm 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 sup- 
posed to bear imperfect flowers, of which the additional 
carpels are the only traces, but this explanation seems 

In addition to the references already cited the follow- 
ing may be given : 

Duchartre, ' Ann. Sc. Nat.,' 4 ser., vii, p. 23 (Tulip). 

Ferrari, ' Hesperides,' pp. 271, 395, 405. Ducbartre, 'Ann. Sc. Nat.,' 

1 ' Linn. Trans.,' t. xxiii, p. 364, tab. 34, fig. 5. 

2 ' Monog. Polygon,' pi. 3, K. f. 12. 

3 ' Flora,' 1856, tab. viii. 

4 Ibid., 1865, tab. ix, f. 6. 

5 ' Adansonia,' vol. iv. 1864, p. 127. 

" 'Ann. Nat. Hist./ 1845, vol. xvi, p. 126. 


i Ber., L844, roL i. p. 294. Maout, 'Lecons Element.,' v.,1. ii. pp. is8-9. 
Aim. Sc. Nat.,' 1865, p. :117 (Citrus Aurcmtw/m). 
Olos, 'Bull. Soc. Bot. Fr.,' yoL xiii ; ' Bev. BibL,' p. 75. Pasquale, 

• Eleddicont Accad. Sc. Fis. e Math. Napoli.' Oc-tr. 1W> (Solanwnl/yco- 


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 hap- 
pens 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. 3-j1). Thus spikelets of 
wheat occasionally produce more than the three florets 
which are proper to them. 1 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. 

' See Schleclitendal, "Bot. Z*jit.." t. xviii, p. 3S1 (Triticum); also 

• Flora/ t. xiv, 1831, p. 5 [Avena). 



A diminution in tlie 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 arortement 
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 dimin- 
ished 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-deve- 
lopment 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. 



Absolute suppression of the main axis is tantamount 
to the non-existence of the plant, so that the terms 
" acaulescent," "acaulosia," &c, must be considered 
relatively only, and must be taken to signify an atro- 
phied 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 depen- 
dent 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 develop- 
ment. 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 projec- 
tions are developed. 1 

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 ease of the rose. 

1 "Theoriede la feuille,' p. 24. 


With reference to this poinl the reader is ref< rred to 
Mr. Bentham's accounl of the morphology and bomo- 

' in the ' Journal of the Linneao 
Society,' vol. x, p. L05. Hoc also write, pp. 71, 77. 

Some doubts also to the nature of the beak 

or columella of such fruits as those of Geramcu 
MahacecB, Urnbelliferce, EuphorUacece, &c. The nature 
of the organ in question may probably be different in 
the several orders named; at any rate the subject can- 
not be discussed in this place, and it is mentioned here, now and then, it happens that th in 

question is completely wanting, and hence affords 
illustration of suppression. 


Tins 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 

The terms aphylly, meiophylly, and meiotaxy may 
be employed, according as the individual leaves are 
altogether wanting, or ace to the diminished 

number of parts in a whorl, or a decrease in the 

Aphylly. — Entire suppression of the leaves is a rare 
phenomenon. Under ordinary circumstances it occurs 
in most Cactaceas, in some of the succulent Buphorbi 

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'- 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 Gliffortia, at other 
times the lateral leaflets are deficient, as in Citrus or 
PhyllartJi/ron. Ononis monopliylla and Fragaria mono- 
phylla 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. 1 

In like manner varieties of the following plants 
occur with simple leaves, Rosa berberifolia (Lowea), 
Rubus Idceus, Robinia pseudacacia, Fraxinus excelsior, 
Sam 1 1) i tc us 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 
Stellatce, and frequently in Paris quad/rifolia, the 
number of leaves in the verticil is reduced. Care must 
be exercised in such instances that an apparent dimi- 
nution arising from a fusion of two or more leaves be 
not confounded with suppression. 

Meiophylly of the calyx or perianth. — A lessened number 
1 'Rev. Hortic.,' 1866, p. 167. 

COROLLA. 'S ( .»/ 

of sepals is not a very common occurrence among 
dicotyledonous plants. Scringe figures a proliferous 
flower of Arabis alpina with two sepals only, and a 
similar occurrence lias been noticed in Dvplotaxis tervwh- 

In Gattleya 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 Cheno- 
pod/iacece, in which the inflorescence is dense, a suppres- 
sion of two or three sepals sometimes occurs. The 
species mentioned are Ambrina ambrosiodes, Chenopodi/u/m 
(jhiucum, and Blitum polymorphum. 

Meiophylly of the corolla. — Suppression of one or more 
petals is of more frequent occurrence than the corres- 
ponding deficiency in the case of the sepals. Among 
GaryophyllacecB imperfection as regards the numerical 
symmetry of the flower is not uncommon, as in species 
of Cerastmm, Sagina, Dianthus, &c. In Barmnculaceos 
the petals are likewise not unfrequently partially or 
wholly suppressed. A familiar illustration of this is 
afforded by Manuncailus auricow/us, in which it is the 
exception to find the corolla perfect. 1 Some varieties 
of Corchorus acutanguhis in west tropical Africa are 
likewise subject to the same peculiarity. Amongst 
Papilionacece absence of the carina or of the aire is not 
uncommon, as in Trifolium r&pens, 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 aire 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." 

1 Be 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 


In Orchidacece entire absence of the labellum, fre- 
quently without any other perceptible change, is of 
common occurrence. The writer has seen numerous 
specimens of the kind in Ophrys apifera and 0. arani- 
fera; also in Dendrobmm nobile, JErides odoratum, 
Oypripedmm villosum, Listera ovata, &c. Morren 1 
mentions analogous deficiencies in Zygopetahim maxil- 
lare, 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, a-x^Xapiov. 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 andrcecium. — 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 Diploiaxis tenuifolia," St. 
Hilaire in Cardamine hirsuta, others in C. sylvatica. 

In CaryophyUaccco suppression of one or more sta- 
mens has been observed in Mollugo cerviana, Arenaria 
tetraquetra, Cerastium, &c. s Among violets the writer 
has observed numerous flowers in which two or three 
stamens were suppressed. Chatin 4 alludes to a similar 
reduction in Tropceolum, while in flowers that are 
usually didynamous absence of two or more of tlie 
stamens is not unfrequent, e. g. in Antirrhinum, Digitalis, 

i 'Bull. Acad. Belg.,' t. xix, part 1, p. 255. 

- 'Bull. Bot.,'i, p. 7, tab. i. f. 7. 

:| See Gay, ' Ann. Sc. Nat..' iii. p. '27. 

4 'Ann. Sc. Nat.,' 4 ser., v. p. 305, 

aiNosciuM. 399 

while in a flower of ( 'ataljpa ;i 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 Trifoliwm r&pens and T. hybrid/urn, and 
lias been seen in Delphinium, &C. 1 

Meiophylly of the gyncDcium. — 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 
Avhich 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 devia- 
tions from the number of carpels that are ordinarily found 
in the particular species under observation. As illus- 
trations the following may be cited : — Arena/da tetra- 
queta, 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 
hi the allied genus Mozlnna. Eeseda luteola occasion- 
ally occurs with two carpels only, while Aconites, Del- 
phiniums, Nigellas, and Paaonies frequently experience 
a like diminution in their pistil. 

In a flower of Pwpaver Rhceas the winter has recently 
met with an ovary with four stigmas and four parietal 
placentas 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 wnld bramble {Embus) 

1 Cramer, ' Bildungsabweich,' p. 90. 


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 Gratcegus the pistil is 
similarly reduced to a single carpel, as in G. monogyna. 

The writer has on more than one occasion met with 
walnuts (Juglans) with a single valve and a single 
suture. 1 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 Oandolle. 

Among monocotyledons Gonvallaria majalis may be 
mentioned as very liable to suffer diminution in the 
number of its carpels, either separately or in association 
with other changes. 2 

Meiopliylly 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 com- 
monly happens, nevertheless, that a defect in one 
series is attended by a corresponding imperfection in 
adjoining ones. Thus trimerous fuchsias and tetrame- 
rous jasmines may frequently be met with, and Turpin 
describes a tetramerous flower of Cobcea scandens. 
Perhaps monocotyledonous plants are more subject to 
this numerical reduction of the parts of several verti- 
cils than are other flowering plants. Thus, in both 
I/ilium lancifolium and L. auratum the writer has fre- 
quently met with pentamerous flowers. In Gonvallaria 
maialis a like deviation not unfrequently occurs. 3 M. 
Delavaud has recorded a similar occurrence in a tulip. 1 

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 

1 See also Clos, ' Bull. Soc. Bot. Fr.,' xiii, p. 96, adnot. 

2 See Cramer, ' Bildungsabweich,' p. 7. Hildebrand, 'Bot. Zeit.,' xx, 
1862, p. 209. 

3 See Hildebrand, ' Bot. Zeit./ xx, 1862, p. 209. 

4 ' Bitll. Soc. Bot. Fr.,' viii, p. 287. 


four stamens were united by their filaments so us i<» 

form two pairs. 

M. Foamier mentions something of the same kind 
in the flower of an Iris. 1 

Orchids seem peculiarly liable to the decrease in the 
number of their floral organs. Prillieux 3 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 Epid&ndrvm Stamfordiamum. In one 
flower of the last-named species the perianth consisted 
of one sepal only, and one lip-like petal placed opposite 
to it. 3 Morren 4 describes a flower of Cyprvpediwrn in- 
signe, 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" (Cypripedivm 
ca/ndidwm) "bears two flowers: the axillary one is 
normal ; the terminal one exhibits the following pecu- 
liarities. 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 actuate 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 

1 ' Bull. Soe. Bot, Fr.,' vol. viii. 1861. p. 152. * Ibid., ix, p. 275. 

3 rbid.. 1861, vol. viii, p. 149. 4 ' Lobelia,' p. 55 




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 

Here, therefore, is a symmetrical and complete, 
regular, but dimerous orchideous flower, the first ver- 
ticil of stamens not antheriferous, the second anthe- 
riferous, 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." 

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 

Fig. 198. — Regular dimerous Fig. 199.— Regular dimerous 

flower of Calanthe vestita. flower of Odontnglossum Alexandre?. 

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 Odontoglossiun Alc.viun.lnr 
a similar reduction of parts 1ms been observed by the 
author (fig. 199). 

It is curious to observe in these flowers how pre- 
cisely one sepal occupies the position of the labclhim, 
and how the lateral petals are displaced from the posi- 
tion 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 Moclenia was established on a malformed 
flower of Orchis of similar character to those above 

Meiotaxy of the calyx, — As already mentioned, this 
term is here employed to denote those illustrations in 
which entire whorls are suppressed. Complete de- 
ficiency 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 Salicinece 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. De- 
ficiency of the corolla was observed in Campanula pev- 
foliata and Buellia clandestina by Linne, who calls such 
blooms flores mutilati. 1 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 Malpighiacece, 
Violacece, OxalidaGece, in some of the flowers of which 
the petals are altogether wanting, while in others the 

1 ' Phil. Bot..'-p. 119. 


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 deve- 
loped 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 de- 
prived of their corolla. 1 

Apetalous flowers have been noted most frequently 
in the following plants : 

Aconitum, sp. pi. ! Crataegus ! 

Cardamine impatiens. Medicago lupulina. 

Cheiranthus Clieiri ! Melilotus officinalis. 

Viola odorata ! Ononis minutissima. 

Cerastium vulgatum ! Saxifraga longifolia. 

Alsine media. Verbascuni Thapsus. 

Stellai'ia. Ajuga iva. 

Lychnis dioica ! Teucrium Botrys. 

Dianthus barbatus, and other Lamiuin purpureum ! 

Caryophyllea?. amplexicaule. 

Helianthemum, sp. ! Polemonium cseruleum. 

Oxalis Acetosella. Campanula, sp. pi. ! 

Balsaniineae. Ruellia clandestina. 

Malpighiacese. Lonicera Periclymenum ! 

Rosa centifolia. Tradescantia, sp. ! 

arvensis ! Hymenocallis. 

The following references apply some to apetalous and others to dimor- 
phic 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 Jussieu, 'Monogr. Malpigh.,' pp. 82,334. Torrey, ' Fl. New 
York,' i, p. 428. Hooker and Thomson, 'Journ. Linn. Soc' ii, p. 7. 
Guillenhn, 'Archiv de Botan.,' i, p. 412. Michalet, 'Bull. Soc. Bot. 
Fr.,' vii, p. 465. Miiller, ' Bot. Zeit,,' 1857, p. 729. ' Natural History 
Review,' July, 1862, p. 235. 

1 ' Cat. Plant. Pyr/ p. 58. 

THE ani)Im:cu;m. M)5 

Meiotaxy of the andrcecium. — 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 
1G35. Cornuti speaks of it in his ' Enchiridion.' In 
1860 M. du Parquet discovered it in peaty woods near 
Nangis (Seine et Marne). 

Many Uinbellifenv, such as Trinia vulgaris, present a 
like deficiency, while it is of common occurrence 
among Bosacece and Pomaccce. In the latter group 
the St. Valery apple, so often referred to, is an illus- 
tration. To obtain fruits from this variety it is neces- 
sary to ajjply 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. 1 
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 

M. Dupont has given a list of nineteen species of 
Chenopodiacece in which female flowers are occasionally 
produced, owing to the entire suppression of the 
staminal whorl. 2 

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 

1 Moquin-Tandon, loc. cit., p. 328. 

: For other instances see Chatin in ' Aim. Se. Nat.,' i ser., vol. v, p. 305 


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 ! ' Trifoliurn hybridum. 

auricomus ! repens. 

bulbosus ! Umbelliferae, sp. pi. 

Cruciferse, sp. pi. Onagracese, sp. pi. 

Violacese, sp. pi. Hippuris vulgaris. 

Honckenya peploides. Callitricbe veiiialis. 
Stellaria. autuninalis. 

Caryopbyllaceaj, sp. pi. Loniceva Periclynienum. 

Malpighiaceae, sp. pi. Erica Tetralix. 

Tropseolurn majus ! Tbymus Serpyllum. 

Fragaria vesca ! Calceolaria. 

Rubus, sp. Composite, sp. pi. 

Pyrus Malus. Chenopodiacese, sp. pi. 

Agrimonia vulgaris. Stratiotes aloides. 
Rosacese, sp. pi. ■ 

Meiotaxy of the gynceeium.— 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 JJmhellifercB, 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 sta- 
mens, 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 Torenla scqbra, accom- 
panied by resorption or disappearance of some parts 

M. ]'», 

and spiral torsion of others. The pistil was entirely 
absent in this instance. 1 

M. Gaetano Licapoli places on record an instance 
where the petals and carpels of Mi lianthus major were 
suppressed. 9 

On the whole, the pistil seems less subject tu cha 
of this character than the andrcecium. 

Suppression of the pistil has been most frequently 
recorded in flowers (normally bisexual) of — 

Ranunculus! Trit'uliuui repens. 
A.conitum '. hybridum. 

Delphinium ! Composite, sp. pi. 

Pa?onia. Datura. 

Cary ophylleae ! Torenia aaiatica. 

UmbeUifc Colchicum aatomnale. 

Suppression of ovules,— abortion of seeds. — The two . 
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 t 
kc. 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, lis when carpels become foliaceous 
and their margins detached. Hybridisation and cross 
fertilisation are also well-known agents in diminish] 
the number and size of seeds. 

1 See also Murren. ' Bull. Acad. Belg.." xv. Fuchsia, p. 67. 

: Cited in ' Bull. See. Bot ., France.* t. xiv (" Rev. Bib!."), p. 253. 



Meiotaxy of the parts of the flower in general. — In the pre- 
ceding sections suppression lias been considered as it 
affected individual members of a whorl or separate 
whorls. It rarely happens, however, that the suppres- 
sion 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 

Maximowicz describes a Stellaria (Krasehenikovia) 
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. 1 

Kirschleger 3 mentions a variety of Lanier ra Ccupri- 
folium, which was not only destitute of petals but of 
stamens also. 

In some species of Muscari and Bellevalia the upper- 
most 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, wag, 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 

1 ' Primit. Flov. Amurens.' p. 57. . 
- ' Flora,' 1848. p. 484. 


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, lli/<trinth/U8 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 Bow-lea volubilis. The 
wig plant (Rlius Cotlnus) affords another illustration of 
the same thing. Some tendrils also owe their appear- 
ance 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 verticil- 
lasters on one side of the stem completely wanting, 
the adjacent leaf being, however, as fully formed as 

General remarks on suppression. — On comparing together 
the various whorls of the flower in reference to sup- 
pression, 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 : — Ranimculacece, Grucif&rce, ( "ari/uphi/llacece, 
Violacece, Legv/mmosce, Onagracece, JasminacecB, Orchi- 
daeecB. 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 

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 
OnagracecBi Oi/rc&ia and Lojjezia may be referred to, 
the former normally dimerous, the latter having only 
one perfect petal. So in fuschias, 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 Violacece, Lecjuminosce, and Orchidacece. 
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 con- 
clusion. This is but another illustration of the fact 
that conditions which are abnormal in one plant con- 
stitute the natural arrangement in others. 

As to the suppressions that occur in the case of 
the sexual organs, and the relations they bear to di- 
morphism, dichnism, &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 tera- 
tology affords very numerous illustrations of those 
intermediate conditions which are also found, under 


natural circumstances, between the absolutely uni- 
sexual 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. 



In the animal kingdom the entire adult organism, as 
well as each of its separate parts, has certain dimen- 
sions, 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. Never- 
theless observation and experience soon show what 
may be termed the average size of each plant, and any 
disproportion between the several organs is speedily 

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 par- 
ticipate in this diminished size, we have what are gene- 
rally 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 maer&phylla, hngifoMa, 
macrantTia, &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 here- 
ditary transmission. It may be supposed that such 
deviations may have originated, in the first instance, 
either from excessive use, or from disuse, or froin 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 varia- 
tions 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 its nit of 


increased growth and those which arise from diminished 
action. It will bo seen, therefore, that in these in- 
stances it is the bulk of the organs that is increased, 
not their number; moreover, their development or 
metamorphosis is not necessarily altered. In connec- 
tion with increased size an alteration of consistence is 
so frequent that the two phenomena arc 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 develop- 
ment. 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. 



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 

' 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 in- 
ordinate development of cellular, fibrous, or ligneous 
tissue, are, of course, strictly homologous with the 
similar changes which occur, under ordinary circum- 
stances, during the ripening of fruits or otherwise. 

Hypertrophy, whatever form it may assume, may be 
so slight as not perceptibly to interfere with the func- 
tions 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 groAvth 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 



A swollen or thickened condition (renflement) is 
usually the result of a disproportionate formation of 
the cellular tissue as contrasted with the woody frame- 
work 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 Eehinocartus, 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 further illustrations of this 
common tendency in some organs to become hyper - 
trophied. 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. 

Enlargement of axile organs. — All the species of Pe- 
largonium, Geranium, Mirabilis, as well as those of 

Fig. 200. — Pelargonium, one branch of which was hypertrophied, 

Qaryophyllece and other orders, have tumid nodes as 
a normal occurrence. In the genus Pelargonium this 
swelling is sometimes not confined to .the nodes, but 

KNAUBS. Il'.» 

extends to the interspaces between them, e. g. I'. 
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 in- 
debted to Dr. Sankey for a branch of Tela/rgonw/m 
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, Di/psacus, &c. (See Spiral 

Knaurs. — On certain trees, such as the oak, the horn- 
beam, 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 irregu- 
larly, 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 in- 
ordinately developed, as if by compensation for the 
curtailment in length. 1 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 

1 On tlie 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. 75b'. 



the "uovoliof 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. 

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 

Fig. 201. — Formation of tubers or hypertrophied buds in the axils of 
1 '.lives in tlie potato. 



becomes apparent. A swollen bud or bulbil in tins 
situation is not uncommon in some cultivated tulips 
mid lilies. The presence of small tubers in the axils 
of the leaves in the potato, as shown in fig. 201, is also 
not unfrequent. 

Enlargement of the flower-stalk. — The cauliflower and 
broccoli afford familiar illustrations of hypertrophy of 
the flower- stalk, accompanied by a corresponding 

Fig. 202.— Inflorescence of ash {Fraximis). with hypertrophied 
pedicels, flowers absent. 



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. 1 

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. 

Fig. 203.— Monstrous pear, showing extension and ramification of the 
succulent floral axis. The bases of the sepals are also succulent. 

1 Jaeger. 'Flora.' 1860 p. 49, tab. i. 



(See fig. 17G, p. 327.) The lower swelling is entirely 
axial in those cases, as no trace of carpels is to be seen. 
M. Carriere 1 mentions an instance wherein from the 
base of one apple projected a second smaller one, desti- 

Fig. 204. — Monstrous pear, showing extension and swelling of axis, &c. 

tute 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. 

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 succu- 
lent 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. ) 2 

' ' Revue Horticole,' 1868, p. 110, figs. 12, 13. 

- The reader may also refer for further information on the subject of 
malformed pears to Ivmisch, ' Flora,' 1858, p. 38, tab. i ; Lindley. ' Theory 


A very curious illustration of hypertrophy of the 
flower-stalk is recorded and figured by M. Carriere 1 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 appear- 
ance was thus that of a crab -apple. There was no 
stone in the interior, but simply a rudimentary kernel 
or seed. 2 

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. 3 

In this place may also be mentioned the hypertro- 
phied 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 un- 
usually large in these flowers, but it also protruded 
beyond the ovary. 4 A similar state of things in Lobelia 
and Cwphea has already been alluded to under the head 
of Alterations of Direction (p. 210). 

The following singular growth in a tomato is dc- 

of Horticulture'; Caspary, ' Bull. Soc. Bot. France,' vol. vi, 1859 (Rev. 
Bibl.), p. 235; Duharnel, ' Phys. Arbr.,' liv. iii, cap. 3, p. 393, fig. 308; 
Bonnet, ' Recherch. Us. feuilles,' tab. xxvi, fig. 2 ; Moquin-Tandon, 
• El. Ter. Yeg.,' p. 384, &c. Some of the cases recorded are, however, 
instances of true prolification. 

1 ' Revue Horticole,' 1868, p. 310. 

- 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 Rosaccce. The case would fitly be included under altera- 
tions of position, but the sheets relating to that subject were printed 
off before the publication of M. Carriere's notice. 

: < ' Bull. Soc. Bot. Prance,' 1860, vol. vii, p. 881. 

4 " Monstr. Veget." in ' Neue Denkschrift." 


scribed by the Rev. M. J. Berkeley in the ' Gardeners' 
Chronicle' for 1806, p. 1217, and appears to have been 
.in extension of the placenta : — " On the first glance it 
seemed as if an unsually large grape-stone had acci- 
dentally 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 compo- 
sition of a portion of the placenta. On making a 
vertical section, instead of being succulent, as I ex- 
pected, 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 placentas or any- 
thing indicating an abortive fruit. On closer exami- 
nation 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 appear- 
ance to them when seen en masse. I am inclined to 
regard the body rather as an abortive axis than an un- 
developed 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 sub- 
stance of leaves takes places in several ways, and affects 
the whole or only certain portions of them. The sim- 
plest 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 in- 
creased 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 bul- 
lata." 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 with- 
out 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 mal- 
formations. But where one part only of the led' 
is excessively developed, the other portion remaining 
in its ordinary condition, there can be no hesitation in 
ranking the phenomenon as teratological. 

Thus, Moqnin sa}\s that the median nerve may be 
prolonged beyond the blade of the leaf in the form of 
;i 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 1 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. 
Uelavaud 2 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 occu- 
pying 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 sup- 
pression ; 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 Campbell). 

1 ,; Sched. de Monst, Plant/" in 'Act. Helvet..' t. ii. pi. ii. f. 14. 
; • Bull. Soc. Bot. France' vol. viii. 1861, p. 144. 



Enlargement of the perianth, &c. — One or all the segments 
of the perianth maybe subjected to hypertrophy; thus, 
the utricle of Gcurex 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 Cruciferce. The perianth of Bwmex 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 SaUola Kali 
and of Chenopodmm mwrale in which some of the seg- 
ments of the perianth were five or six times larger than 
they should be. 

The adjoining woodcut represents a singular con- 
dition of some cocoa-nuts in the Kew Museum, the 

Fig. 205. — Hypertrophy of the perianth in Corns nucifera. 


appearance of which is due apparently to an hypertro- 
phied condition of fclie 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 in- 
creased to such an extent that the fruit appears sur- 
rounded 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 Gin- 
chonacece, e.g. Musscenda, Pinchneya, Galycophylhm, 
one or more of the calycine lobes are normally dilated 
and petaloid, the others remaining small and compara- 
tively inconspicuous. Inequality in size is, indeed, a 
common occurrence in the sepals of many natural 
orders — Polygalacece, Legummosce, Ldbiatce, &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 Pa/paver Rkosas, Reichenbach in 
Campcmula persicifolia, and A. de Candolle in G. Ba- 
pnncalus. M. Brongniart also has recorded 1 a remark- 
able variety of Prwmla sinensis cultivated in the Jardin 
des Plant es at Paris, wherein the calyx is enormously 
developed. MM. Founder and Bonnet have described 
flowers of B/ubus with hypertrophied calyx in conjunc- 
tion with atrophy and virescence of the petals and 
other changes. 2 

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 
TKpsacus, and also mentions a remarkable variety of 
Viola odorata cultivated in the neighbourhood of 

1 'Ann. Sc. Nat..' ser. 2. t. i. p. 308, pi. ixc. fig. 1. 
- 'Bull. Soc. Bot. France,' 1862, t. ix. p. 37. 


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 1 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 Solarium Dulcamara in which 
all the upper flowers had two or three stamens of larger 
dimensions than the others. This happens habitually 
in Solatium 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 Solanacece and the Scrophu- 

Enlargement of the gyncecium. — In some flowers which 
have become accidentally female the pistil becomes 
unusually large, and even to such an extent as to pre- 
vent the passage of the pollen. Moquin remarks having 
seen this enlargement in the pistils of Suceda 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 3 have figured some anomalous 
flowers of Diplotaxis tenuifolia in which the pistil, more 
or less distended and deformed, was considerably elon- 
gated below, so that it seemed to be borne upon a long- 
stalk, analogous to that of fruits of Capparids. Dr. 
Khnsman 3 mentions an instance of a similar kind com- 

1 • Consid. org. Fleur.,' Montpell, 1829, 25, 26, pi. ii, f. 18 and 19. 
- ' Bull. Bot.,' t. i, p. 7, tab. 1. 
;i ' Linnsea,' vol. x, p. 604, tab. 5. 


bincd with hypertrophy of the Bepala and pistils; in- 
deed, the alteration is not uncommon among Crucifers. 
Pyrethrv/m inodorwm is very subject to hypertrophy. 
The styles of its radial florets become elongated with- 
out any other alteration ; at the same time the small 
corollas become green, and show a tendency to assume 
a foliaceous condition. Sometimes the hypertropliv 
affects also the styles of the central florets, and thesi 
also become enlarged to double or treble their usual 

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 1 records an instance in 3umea 
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 
a/rticulatus is thus punctured, it acquires a size two or 
three times larger than ordinary, becoming at the 
same time sterile. 2 

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 Sinnmgia 
yurpwrea 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. 3 

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 

1 ' Mem. Acad. Sc. Toiiloxise,' 5 ser., vol. iii. 

' ; • Re. nosol. Veget.,' pp. 342. 

3 • IUustr. Hortio./ 1868, Misc.. p. 62. 


of pea with pods double the ordinary size. 1 M. Clos 2 
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 Legwrdnosce, as Medicago lupulina, Meli- 
lotus leucantha, the carpels are sometimes hypertrophied 
and elongated, so as to resemble a claw or hook. 3 

The fruit of the common groundsel (Senecio vul- 
garis) 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 be- 
come 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 Orinum, Amaryllis, and Agave. 
These are true seeds enormously dilated. 4 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 circum- 
stances. 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 

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 suc- 
culent than ordinary, from the formation of an in- 
ordinate amount of loose cellular tissue. Generally 

1 ' Ann. Soc. Linn.,' Paris, t. i, p. 139. 

2 ' Mem. Acad. Toulouse,' t. 6, 1862. 

3 ' D. C. Prod.,' ii, pp. 172, 187. 

* Richard, " Obs. sur les bulbilles des Orinum;" 'Ann. Sc. Nat.,' t. ii, 
p. 12. pi. i, fig. 1, 2. See also A. Braun, "Memoire sur les graines 
charnues des Amaryllidees," &e. ; ' Ann. Sc. Nat.,' 1800, vol. xiv, p. 1, 
tab. 1. 


speaking, bhe appearances presented in such cases are 
not sufficiently striking to demand notice other than 
as regards their size One illustration, Ik > we vcr, may 
bo 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. 



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 elon- 
gation that takes place in plants wdien 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 planta- 
tions, 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 r in running water their roots, stems, and some- 
times their leaves, become excessively elongated, as in 
Rammculus 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 Pota/mogeton bifolium, a plant which Mr. 
Bentham subsequently discovered to be nothing but a 
flowerless variety of Vicia Faba distorted by its growth 
in water. 1 

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 lengthen- 
ing 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 
Umbelliferce the umbels are occasionally met with sup- 
ported 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 Legimdnosce, Trifoliwm r&pens, 
&c. &c. 

Another illustration of the sort is that recorded by 
M. Founder, wherein the usually umbellate inflorescence 
of Pelargonium was, through the lengthening of the 
main stalk, transformed into a raceme. Among Com- 
posites a similar change may sometimes be met with. 

1 -Cat. Plant." Lang., p. 113 

I l.<>\\ BB-8TALKS. 1"> ; < 

MM. Clos and De Schonefeld have recorded the ex- 
istence of a variety of the sweet chestnut (Castanea) in 
which tlir female catkin.- were as Long, and bore oearlj 
as many flowers, as the male spikes. This i- 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 Been bearing thirty 
to thirty-five large nuts. 1 

In the strobile of the hop, Humufois Lwpulus, 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 extru- 
sion or eversion of an ordinarily concave axis, is the 
fig described by Zuccarini," and from the appearan 
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 ac- 
companied 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 Amentacece. (See p. 204, figs. 105, 106.) 

A leng;thenm°; of the axis of the female strobiles of 

DO w 

< 'onifercB is not of infrequent occurrence in Gryptorm na 
ia/ponica, Larix europcea, &c, and this is usually asso- 
ciated, as has been before stated, with a leaf-like con- 
dition 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 elon°'ation of the main rachis has 

O m t 

been considered. A corresponding deviation occurs u. 

1 -Bull. Soc. Bot. France, 1 i. i. L854, p. 1.73, and t. xiii. p. 96. 
- Abhandl. Math. Vhy I ' Band, iv, Abhandl. i. tab. i. 



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 re- 

Fig. 206. — Inflorescence of Ranunculus acris, with secondary peduncles 

presents 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 Trifoliwm r&pens, Lotus corni- 
culatus, &c, what is usually a compacjt spike, or head 

LEAVES. 1)37 

of flowers, becomes a raceme from the elongation of 
the pedicels. In Umbellif&rcB 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 Garni, 
Thysseliwwm palusi/re. 1 In Com/positce, when affected 
by an analogous change, the capitulum assumes the 
appearance of a simple umbel, as in Hypochceris r<i<Hc<ii<<, 
Senecio vulga/ris, and other plants. 

In some of the double-flowered apples which have 
been previously alluded to, the flower-stalk is inordi- 
nately 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 repre- 
sented 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, c. 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 Umbelliferce and Com- 
positce, changing very materially the general aspect of 
the inflorescence. Occasionally, also, the leaf-lobes of 
parsley (Apium Petrosclinum) and other crested-leaved 
plants may be observed to lose their ordinary wavy 
form, and to be lengthened into flat riband-like seg- 
ments, 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 lengthen- 
ing of the midrib or central stalk, convert a digitate 
or palmate leaf into a pinnate one. In these instances 

1 See Cramer, ' Bildungsabweich,' pp. 62—79, and Fleischer, ' Missbild. 
der Cultnrpflanzen.' 



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 

Fig. 207. — Portion of leaf of parsley, showing the change from short 
wavy to long flat leaf-lohes. 

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 

In the white clover, Trifolium repens, a similar tran- 
sition may often be observed, as also in some species 
of Potent ill a. 1 

Elongation of the parts of the flower. — The only circum- 
stance that needs especial mention under this section 
is the great lengthening that sometimes takes place in 

1 Schlechtendal, 'Bot. Zeit.,' 1844. p. 457; ' LinnEea,' xi. p. 301, xiv. 
p. 363 ; 'Bot. Zeit.,' 1856, p. 72 ; Masters, ' Rep. Brit. Assoc./ Manchester. 
1861 ; Coultas, ' What may be learnt from ;i tree,' p. 118. 



the carpels, Bometimes as a resull of injury from in- 
Becfca or ftuagufl, a1 other times withoul assignable 


Fig. 208.— Leaves of horse-chestnut, Msculus, showing passage from 
"digitate to pinnate leaves. 

In the case of inferior ovaries this lengthening is, 
perhaps, even more common, as in Umbellifer(B, Gom- 
positce, &c. The common groundsel (Senecio mtya 

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. 1 

Apostasis. — Engelmann made use of this term to ex- 
press the separation of parts one from another by the 
unusual elongation of the internodes. 2 He drew a 
distinction between the separation of individual organs 
one from the other, and the corresponding displace- 
ment 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 Passi- 
florece, Caryophyllece, Capparidece, &c. &c. 

A similar elongation of the thalamus, bringing about 
the separation of the floral whorls, or of their con- 
stituent 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, 

1 For further details refer to the chapter on Displacements, p. 86. 

2 ' Dc Antholysi,' p. 42, § 49. 


i n 

and are, in some cases, united bytheir margins, so that 
a spiral sheet or tube is formed, surronnding the axis. 
This frequently occurs in double flowers of the Ghini 
primrose, I' rim "In sinensis. 

Engelmann 1 figures a case wherein the calyx of 
Anagallis yhcenicea was separated by a rather long in- 
terim* le from the corolla, and a like illustration in 
Torilis Anthriscus. 

Fig. 209. — Flower of Delphinium, showing apostasis of carpels, from 
lengthening of the thalamus, &c. 'Cramer.) 

1 Loc. cit., tab. 2. f. 6. 


A frequent change in Crucifers is due to the forma- 
tion of a long stalk bearing the pod, and thus giving 
rise to the appearance met as a constant occurrence in 

In Tropoeolum 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 pro- 
lification) . 

One of the best-marked illustrations of these changes 
occurs in a permanent malformation of Epilobium hir- 
sutum, specimens of which were originally obtained 
froin 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 men- 
tioned, cites the following plants as having manifested 
this change : 

*Convallaria majalis ! , % Tulipa Gesneriana .'. Veronica 
Ghamwdrys, Orobanche gracilis, Solanwm I/ycopersicum, 
Gentiana campestris, Hypericum, Helleborus fetidus, 
Galiha palustris, Brassica oleracea! and many Rosacea . 
Garyophyllea^, Cruciferce, and Banunculaceo3. (See 
Dialysis, Median Prolification, &c.) 

Apostasis of the subfloral or involucral leaves is not 
of infrequent occurrence in malformations affecting 
Gompositce and Umbelliferce. In the following genera 
it has been observed with especial frequency : — Torilis 
Anthriscus, Eryngium, Athamanta Cervaria, Leontodon, 
Tragopogcm pratense!, Wedelia perfoliata! In garden 
anemones, also, it is a common deviation. 



Ondeb 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 repe- 
tition or to a partition of any organ, but are out- 
growths from others previously formed. 

In prolification and in multiplication the adven- 
titious 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 in a plane 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 project- 
ing from petals, as in Ca/ryophyllece, Sa/pmdacece, &c, 
the coronal filaments of passion-flowers, the cup of 
Narcissus, the appendages that beset the segments of 
the perianth in Lilivm lancifolimn, and other similar 
growths, may be referred to a like process. In many 
cases this has been proved by a study of the develop- 
ment 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 develop- 
ment 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 struc- 
ture, 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 sup- 
posed, since it is well ascertained that, in some cases, 
the formation of the parts of the flower, e.g. the sta- 
mens 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, occasion- 
ally met with in trees, and which are more closely con- 
nected 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 Legu- 
wAnosce, nor of the peculiar excrescences on the roots 
of Taxodi/um distiehmi, as these appear to be normal 
formations. But it may be well to mention in this 

lA \TKlX. 

I 1 ,5 

place an anomalous development which occurs occasion- 
ally in B/U8CU8 aculeatus, and in which, from the upper 
surface of the ordinary flattened leaf-like branch, pro- 
jects, at right angles a second similar branch, so that 
in section the appearance would be like thai of the in- 
verted letter t ; thus, X- 

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 lamina? projecting at right angles from 
the primary one may frequently be observed. In the 
instance figured (fig. 210) the new growths proceeded 

Fig. 210. —Section through base of midrib of cabbage leaf, showing 
supplementary laminae, &c. 

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. 

The development of secondary leaves from the sur- 
faces 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 J2^. In an orange leaf presenting this ap- 
pearance 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. 1 (See p. 33.) 

1 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 whicb 
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 lamina?, 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 ab- 
normal 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 : \0 0/. The nature of the case may be even better seen 

by comparison with the normal arrangement, which would be q . 

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 : 

,0 i , the X in all cases representing the position of the axis, the 

O that of the axillary bud, and the — that of the. lamina?. 


I 17 

Such loaves as those of the hedgehog holly, Ilex 
Aquifoliv/m, var. ferox, and, to a less extent, bullatc 
Leaves, may also be mentioned here as illustrations <>f 
hypertrophy or enation. 

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 mar- 
gin of the leaves, especially, the result is a wavy or 
crisped appearance, " folia undulata, vel crispa." 1 These 

1 Linn., ' Phil. Bot.,' § 274. The term " crispa " is surely preferable 
to that of Re. " phyllorhyseme." 


conditions occur normally in such leaves as those of 
Bumex crisjms, 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. 1 

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 Caryophylleai, Sajrindacece, &c. &c, may be ex- 
plained, as already remarked, by this process, rather 
than by fission, chorisis, or by substitution of petals 
for stamens, &c. Each case must, however, be ex- 
amined 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 

1 See C. Morren, " Consid. sur les deformations," &c, in ' Bull. Acad. 
Belg.,' 1852, torn, xix, part 3, p. 444 ; and as to ferns, see Moore. 
' Nature-Printed British Ferns,' 8vo ed., where numerous illustrations 
are given. 


of alternation) thai suppression of certain organs has 

taken place. 

Taking first those instances in which the supple- 
mentary petals appear on the inner surface of the 
corolla, as being at once the most frequent, and as 
presenting the closest analogy, with similar conforma- 
tions, under natural circumstances, certain double- 
flowered varieties of the Chinese primrose, Primula 
smeTisiSi 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 fimbri- 
ated 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 supple- 
mentary 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 supple- 
mentary petal is turned towards the inner or upper 
surface of the primary corolla, thus ^Z^ ; while, in other 
instances, the front of the adventitious lobe is directed 
towards the corresponding surface of the original petal, 
thus CZ^. Whether these supernumerary petals are 
formed by chorisis or by enation cannot, with cer- 
tainty, be determined without examining the early 
stages of development. 

Of more interest are those instances where the ad- 
ventitious 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 

Fig. 212. — Datura fastuosa. True corolla turned back to show tlie 
supernumerary corolla with the petal-like segments attached to its 
outer surface (reduced). 

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 
























The X indicating the position of the petalodes. 

A still more singular case is that of a variety of the 
Gloxinia, described originally by Professor Edouard 
Morren, 1 but which is now becoming common in 
English gardens. When first observed these flowers 

Fig. 213.— Gloxinia, with supernumerary segments on the outside of 
the true corolla. 

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. Beinp; 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. 

1 ' Bull. Acad. Belg.,' t. xix, p. 224, tab. i ; and ' Gardeners' Chronicle,' 
1865. p. B65. 


Professor Morren considers the adventitious peta- 
lodes 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 suppres- 
sion of all the parts of the flower, except the lobes in 

Fig. 214. — " Catacorolla" of Gloxinia, formed from the union of ad- 
ventitious 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 con- 
fluent, 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 adven- 

I BOM 'I m: I ABPELS. \ : >''< 

tit ions lobes from the outside of the minute partially 
developed petals could be distinctly seen. 

Enation from the stamens. — An illustration of this pr< 
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 pseudocorolla. 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 

^ — ~v. This disposition was well seen in the anther, 
00 ! 

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, 
gfoino- from within outwards, the cut edo;es of two 
perfect polliniferous lobes m the centre ; and on either 
side the petaloid wing representing the remaining 
anther-lobe ; outside these were the edges of the re- 
maining 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 extra- 
ordinary cornute oranges described and figured by 
Ferrari, Gallesio, and other winters on the genus Citrus, 
may be mentioned under this head. A similar forma- 
tion occurs in the fruit of some species of Solarwm. 
(See p. 316.) 



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 struc- 
ture is so incomplete that the office cannot be per- 
formed. These differences depend, of course, upon 
the stage of development which the organ had reached 
when its growth was checked. For practical pur- 
poses 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 circum- 
stances, 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. name), or those in which the leaves or flowers are 
smaller than usual (var. pa/rvifoMce, v. parvifiorce), are 
truly regarded as variations, and not as malformations 
properly so called. 

ABORTION, ]■>'> 

At rpphy is partial and special in its operation, nanism 
is general. 

Cinder ordinary circumstances atrophy is exemplified 
by the presence of rudimentary or imperfect organs, as, 
for instance, in Pentstemon s SwopJmlaria, &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. 



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 ar- 
rested in its growth, the habit and general appearance 
of the plant are materially altered, as in the so-called 
stemless plants, plcmtce acaules. In these the inter- 


nodes are so slightly developed that the leaves are 
closely crowded in tufts or rosettes. When this short- 
ening of the stem (acanlosia) 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 1 cites an instance of Ga/m/pho- 
rosma monsjjeliaca, 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 Artdbotrys ; 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 be 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. 2 

In the birch an arrest of development in some of the 
branches is of common occurrence. The branch sud- 
denly 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. 8 

1 ' El. Ter. Veg.,' p. 132. 

2 Spinosce arbores cultura scepins deponunt spinas in hortis, ' Linn. Phil. 
Bot.,' § 272. 

;i Mr. Sclby. in bis ' History of British Forest Trees,' p. 465, gives 


A shortened condition of the flower-stalks occurs 

occasionally, greatly altering the general character of 
the inflorescence. This has been observed in pelar- 
goniums and in the Chinese primrose, in both of which 
the effect was to replace the umbellate form of inflor- 
escence 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 Gompositce and TJmbelliferee. In the former 
natural order the following plants have been met 
with in this condition : — *Tragopogon pratense /, 
*Cirsium arvense, Hypochceris radicata, Senecio vulgaris I ', 
Coreopsis Drummondi. In the latter order, Daucus 
Garota! CEhianthe crocata! and Thysselinum palustre, 
seem most frequently to have been observed in this 
state. 1 In some gourds the receptacle may be seen 

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 increas- 
ing 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 

1 See Cramer, 'Bildungsabweich.,' pp. 53, 64, for further references. 


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. 

JMoquin 1 mentions having seen the leaves of Glieno- 
podium vulvaria, and of Dvplotaxis muralls 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 polymdrphum. 

Nicandra ph/ysaloides* 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 monstro- 
sities, as in the case of the entire-leaved varieties of 
those plants which ordinarily have cut or divided 
leaves, e.g. Plantago Coronojpus, var. integrifolia, Palaver 
Rlioeas mtegrifolia, &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 aqaritica, the spiney leaves of Berbe- 
ris, 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 malfor- 
mation. 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 

1 'El. Ter. Veg.,'p. 124. 

c Schlechtendal, ' Bol. Zoit..* 1*57, vol. xv. p. 67. 

i m: i.i w ES. 459 

thus altered, but the following species appear to be 
particularly subject to this change : ffisoulus Hippocas- 
t'litiini, Digitalis purpurea, Moras alba, Vagus silvaMca 
contraQta (hort.), Codicewm variegabwm var. erosum 
(hort.), Broussonettia papyrifera, Scolopendriwm mlg< 

Frequently this irregular diminution in proportion 
is coexistent with an unusual degree of cleavage or 
laciniation of the margin, as in Acer plata/noides lacirria- 
bwm, Tilia asplenifolia } Akius 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 lamina? one with the 
other. This has been observed amongst other plants 
in Veronica latifolia, Broussonettia papyrifera, Godiam/m 
variegatwm var. mterruptum (hort.), Scolopend/rium mil- 
gave, &c l (See p. 328.) 

In some of the leaves which have been already re- 
ferred 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 bul- 
lata." In illustration may be cited various species of 
Mentha, Perilla, Ooleus, Fagus sUvaUca crispa, Gytisus, 
Laburnum var., and other forms, cultivated in gardens 
for their singularity. 

Entire absence of the stalk of the leaf occurs nor- 
mally in sessile leaves ; on the other hand the blade of 
the leaf is only occasionally developed in the phyllodi- 
neous Acacias, in some species of Ore///.*, Indigofera, 
Lebechia, Ranunculus, Bupleurum, &c. 

1 On the subject of this paragraph the reader may consult A. Braun. 
"Ueber abnorme Blattbildung," Ac., in * Verhandl.,' d. 35. 'Naturfor- 
seherversamnilvtng ;' Jaeger. 'Flora.' 1850, p. 481, tab, 1. Digitalis. 


De Candolle, 1 from a consideration of Strelitzia jnncea, 
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 inner- 

FiCr. 215. — Inner leaves of cabbage reduced to their midribs. 

most 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. 

The suppression of one or more leaflets of a com- 
pound 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 

1 ' Org. Veget.,' i. p. 286. 


extremities they bear crowds of imperfect flowers, in 
which the calyx only is visible, and that only in a rudi- 
mentary 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 

Fig. 216. — Abortion of four out of five petals, Viola tricolor, side and 
front views. 

that of the calyx. In Armaria serpyllifolia the petals, 
especially in autumn, are only one fourth the length of 
the sepals. Anagallis phoenicea, HoncJcenya peploides, 
Arabis alpina, Ranunculus auricomus, Rubus fruticosus, 
and Geranium columbinum, also frequently afford illus- 
trations of this circumstance. 

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 habit- 
ually among Violacece. 

The diminished size of the petals sometimes coexists 
with an increase in their number, as in a flower of 
Streptocarpus Rexii, mentioned by Bureau. 1 

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 

1 ' Bull. Soc. Bot. France,' vol. viii, 1861, p. 7L0. 



p. 347, fig. 179), a few less common illustrations may 
be cited. In crocuses it is not a very uncommon cir- 
cumstance 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 1 mentions a flower of Fritillaria im- 
perialis 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 2 records a similar 
occurrence in the same plant. 

Morren 3 gives details of like appearances in Hymeno- 
callis americana, and Delavaud 1 in Tigridia jpavonia. 

In certain orchids an arrested development of the 
perianth is habitual, as in Oncidium abortivum (fig. 
217), where, on a large branching panicle, numerous 

Fig. 217. — Flower of Oncidium abortivum, magnified. 

abortive, but few perfect, flowers are produced. In a 
similar way the petals and labellum of Odontoglossu/m 

1 ' Linnsea,' 1830, vol. v, p. 492. 

2 ' Mus. Senkenb.,' ii. p. 45. 

:i ' Bull. Acad. Roy. Belg.,' 1851, t. xviii, part i, p. 27. r ,. 
4 ' Bull. Soc. Bot. France,' vol. viii, 1861, p. 147. 

9,1 LMENS. t63 

Uro-Ski inter i have been found reduced to filamentous 


Abortion of the stamens. — Atrophy of one or more sta- 
mens is of very common occurrence, as a general rule, 
in many genera of plants, e.g. Scroph/ularia, Erodi/um, 
many Eestiacece, &c. &c. As a strictly teratological 
condition atrophy of the stamens is more rare than 
complete suppression. It has been noticed in Arabis 
(i//>iiiii, Gerastvu/m ghmeratum, C. tetrandrv/m 3 Rha/mnus 
cathartic us. Anemone, Hepatica, &c. It happens fre- 
quently 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 sta- 
men 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. 1 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 appa- 
rently perfect, even to the ovules, the stamens were 

1 See Darwin, ' Variation of Domest. Aniin. and Plants.' ii. 165. 



atrophied, and distorted, and contained little or no 
pollen ; the few grains of the latter being smaller than 
nsual. (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. 
StercuUacece, Eupliorhiacece, Bestiacece, &c. &c, and in 
some natural orders there appears to be a tendency 
towards a dioecious condition, e. g. Oaryophyllece, 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. 1 The 
ovary of aconites, according to Moquin, is very subject 
to atrophy. 

Fig. 218.— Bladder plum. 

During the maturation of the pistil, and its passage 
to the fruit, great changes of consistence frequently 

1 Gay, ' Ann. Sc. Nat.,' ser. i, 1824, t. iii, p. 44. 

pistil. 465 

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. 1 

MM. Fournier and Bonnet 2 describe a fruit of a 
Rulms, with perfectly dry fruits, like those of a Qewm, 
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. 3 

Schlechtendal describes 4 the ordinarily baccate fruit 

1 See De Candolle, ' Mem. Legtim.,' tab. 3, f. 1 ; Wyville Thomson, 
' Trans. Bot. Soc. Eclinb.,' 1851, July 10th ; Berkeley, ' Gardeners' Chro- 
nicle,' 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 Bruns- 
wick, 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. 

- ' Bull. Soc. Bot. Fr.,' 1862, vol. ix, pp. 37 et 291. 

3 Carl Schimp, ' Fl. Friburg,' vii, p. 745 ; Hook, fil., ' Journ. Linn. 
Soc.,' vi, p. 9. 

4 ' Linnsea,' vol. v, 1830, p. 493. 



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, leav- 
ing 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 flat- 
tened one. 1 See also under Heterogamy, Meiophylly, 

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. 2 This is the case, under ordinary circum- 
stances, 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 ana- 
tropal, 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 

1 Moquin-Tandon, 'El. Ter. Veg.,' p. 325. 

2 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. 


or imperfect development of the indusium, as in what 
are termed exindusiate varieti 

General remarks on abortion, coincident changes, &c. — Refer- 
ence has already been made, while treating of hyper- 
trophy, suppression, Arc., to certain other char, 
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-hya- 
cinth, Muscari c&mosum, var., monstro&um, the absence 
of flowers is compensated for by the inordinate for- 
mation of brightly coloured threads which appear to 
be modified pedicels pp. 347, 348) ; so also in the 

wig plant, Rhus Coin So the atrophy of the sta- 

mens, in some flowers, is coincident with the hyper- 
trophy 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 Desmodi/u/m marylandicu/m were atrophied, 
while the calyx and legume, on the other hand, were 

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 2 
on the different degrees of atrophy up to complete sup- 
pression, observable in the flowers of BeUeoaMa 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, 

1 See Moore, ' Xature-Printed Ferns,' 8vo, for numerous illustrations 
both of depauperate and exindusiate ferns. Scolopendrium vulgarc 
seems to be one of the ferns most commonly affected in this way. 
:-. loc. cit., vol. ii, pp. 135, 147. 159. loo. &c 

- ' Bull. Acad. Belg.,' t. xvii, p. 3?.. t. 1 ; Lobelia, p. &5. 


the filaments often being developed without anthers. 
Ultimately a deformed and empty perianth alone re- 
mains. In the ordinary course of things the mouth 
of the perianth is open, but in some of these malfor- 
mations it is closed, and when that happens, the effects 
of atrophy are the more observable in the stamens and 

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 Beccabwnga, 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 may be 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 1 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 con- 
siderable 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 hetero- 
geneous 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. pe- 

1 Cited in ' Henfrey's Botanical Gazette,' i, p. 179. 


culiar 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 
iu the case of Arenaria tetraquetfa, which is polygamous 
when growing in mountain districts. Other illustra- 
tions of a similar character are mentioned under the 
head of Heterogamy (p. 19G). 

Pressure has been already alluded to as one of the 
most obvious of the inducing causes of atrophy and 

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 relat- 
ing 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 cer- 
tain that internal causes inherent to the organization 
of the plant are more important. Mr. Darwin 1 accounts 
for the existence of rudimentary organs by the opera- 
tion 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 hermaphro- 
dite flowers tend to become dioecious, as in Ccuryojphyl- 
lacecB, Orchidacece, Plantaginacece, Primulacece and other 

1 ' Origin of Species,' p. 450. 


Although many of the circumstances above men- 
tioned 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. Yeget.,' p. 120 ; 0. Morren, " De l'atrophie en general," in 
' Bull. Acad. Belg.,' t. xyiii, 1851, part i, p. 275. 



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 under- 
stood 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 perver- 
sion of development. 

Under natural, i. e. habitual circumstances, the for- 
mation 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 demarca- 
tion can be drawn between cases of perversion and of 
arrest of development. 

Formation of scales. — These may be mere epidermal ex- 
crescences, 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 perulas of leaf-buds, or the paleas on the 
common receptacle of composite flowers. Other illus- 
trations of a like character are to be met with in the 


membranous scales that represent leaves in Buscus, 
Aspa/ragus, Pw/us, &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. Sa/pvndacece 3 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 i 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 sepa- 
ration 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 occa- 
sional presence of the perfect in place of the rudimen- 
tary 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 Vic la, 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 fox- 
glove 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 repre- 
sent 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 

Formation of hairs, spines, &e. — The adventitious produc- 
tion 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 de- 
velopment 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 Pa/rowychia 

s&rpi/l/ij'olia. 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 morpho- 
logical significance of these structures. Thus, in 
Passiflora and Viburnum Opuhis, 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 hypo- 
gynous disc. 1 Moquin has seen similar instances in 
the flowers of a Rose, Hypericum, and Poppy. 

M. Planchon 2 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. Gron- 
land and Trecul. 3 

Dr. Hooker shows that the pitcher of Nepenthes is 
due to a modification of a gland placed at the extremity 
of the midrib. 4 

Formation of tendrils. — These are of very varied mor- 
phological 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 

1 ' Consid. Org. Fleur.,' p. 44, pi. ii, fig. 23. 

2 'Ann. Sc. Nat.,' 3 ser., Bot. ix, pi. 6, ff. 1, 2. 

3 ' Ann. Sc. Nat.,' 3,ser., Bot. 1855, pp. 297, et 303. 

4 'Trans. Linn. Soc.,' xxii, p. 415. 


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 gene- 
rally belongs more to general morphology than to 
teratology. 1 

Kirschleger, however, has recorded the existence of 
a cirrhose sepal in Curcurbita Pepo." 

1 See Darwin, " On Climbing Plants," ' Journal of Linnean Society,' 
vol. ix, p. 1. 

3 ' Flora,' 1845, p. 615. 


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 dis- 
turbance 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 Candolle, 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 sub- 
ject of tli>' development and construction of the leaf 
further than they are illustrated by ordinary terato- 
logics] phenomena. 

From this point of view perhaps the most interesting 

circumstance is the part that the sheath of the leaf 
plays. 1 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, corre- 
spond 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 con- 
nected 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 Candolle to consider the 
entire leaf as a composite structure. The morpho- 
logical unit, says he, is the cellular protrusion or 
growing point (saillie) and its corresponding fibro- 
vascular bundle. 2 

The identity, in a morphological point of view, of 
the leaves and the lateral parts of the flower is so 

1 See Clos., ' Bull. Soc. Bot. Fr.,' 1856, vol. iii, p. 679. 
; ■ Theorie de la Feuille,' p. 26. 


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 develojDment of ovules on the edges of 
the anther, the co-existence of pollen with ovules on an 
antheroid body, and, stranger still, the actual develop- 
ment of pollen within the tissues of the ovule itself! 
From such facts, in addition to what Ave 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, morpho- 
logically 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, maybe 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. ATe 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 ; l 
see pp. 174, 177, 445, &c. The structure that we 

1 An additional illustration of this may be cited., which has been 


are apt to associate exclusively with one is found to 
pertain to the other. The arrangement of tl ilar 

cords in the leaf-organ finds its counterpart in the axis, 
generally, it is true, modified to suit altered circum- 
stances or diverse purposes. In some cases the dis- 
position is absolutely indistinguishable in the I 
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, morpho- 
logically 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 undifferen- 
tiated state in the form of a thallus among Cryptogam-. 
but which in the higher groups of plants becomes 
marked out into separate portions, each portion hav 
its own distinct functions to fulfil for the common 
benefit of the whole organisation. 1 

Special morphology. — Under this heading brief reference 
may be made to some of the organs whose morpho- 
logical 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 de- 
partments 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. 

brought under the notice of the writer by Dr. "Welwitsch reeen 
in -which some of the leaflets of the pinnate leaf of a species of Mi 
labium were absent, and their place supplied by flowers arranged in 

1 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 Xepaul 
barley, p. 174. 


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 insepa- 
rable from the ovary. The question morphology has 
to solve is whether this tubular structure is to be con- 
sidered as a portion of the axis, or whether it is to be 
regarded as composed of the confluent bases of the 

Mr. Bentham, who has recently reviewed the evidence 
as to the nature of the calyx-tube in his paper on 
Myrtacece, 1 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 con- 
clusions upon such facts as the following : the circum- 
stance 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 andrcecium 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 diver- 
gence of the stamens as part of the receptacle. A 
similar remark applies to the carpels and pla- 
centas. Mr. Bentham further considers that the 
gradual disconnection of the various whorls, that may 
be traced in many plants, is a further proof of concre- 
tion, 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. 3 

Organs originally free and distinct become ultimately 
combined at the base by the gradual protrusion from 

1 ' Journ. Linn. Soc.,' vol. x, p. 103 et seq. 

2 See also the veceptacular tube (ovary P) of Baeclcea bearing stamens, 
see p. 183. It would be natural to see stamens springing from the recep- 
tacle but not from the ovary. 


fehe receptacle of a ring or tube under them, as in the 
stamens of Legv/mmosce; yet, says Mr. Bentham, no 

one would propose to describe the stamina! tube of 
monadelphous Legv/mmosoe 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 Pedicu- 
laris, 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 Rosacece, Myrtacecr, Melasto- 
macece, Passiflora, 1 &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 por- 
tion of the receptacle. 3 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 Tropoeolum or Nebu/mbium 
become very significant. In both the leaf-stalk is 
cylindrical and traversed, as in the case of all cylin- 

1 In Passiflora tlie organogeny of the flower clearly shows the truth 
of this assertion, as was indeed shown by Payer and Schleiden. 

2 See Payer, ' Organ. Veget.' 



drical 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, pre- 
cisely 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 Can- 
dolle 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 mor- 
phological 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, 1 be 
upper part of the carpels in these fruits projecting 
ond tl 10 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 1 lie 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 com- 
pared the production of ovules on them to the forma- 
tion of buds on the leaf of BryophjUnm. Others 
have been led to see in each placenta, even when 
it is, to all outward appearance, a portion of the car- 
pellary leaf, a direct prolongation from the axis, ad- 
herent to the leaf. Teratology shows that ovules may 
be formed in differently 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, 1 there is surely little ground 
for considering the placentas, or o villiferous portions 
of the plant, to be of necessity axial. Here again, 
much of the difficulty vanishes if the morphological 

1 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 iinpari-pinnate leaves 
in which the terminal leaflet is jointed to the common rachis, and in 
the leaves of some Meliacece which continue to push forth new leaflets 
even after the leaf has attained maturity. 


identity of the leaf-form and of the stem-form be ad- 

Structure of the ovale. — 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 noriferous 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 tera- 

Morphology of conifers. — The nature of the pseudo-leaves 
of Sciadcrpitys, and probably of other Conifers, is illus- 
trated 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 con- 
sidered, 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 
Beparate in the adult state, lias been traced to an 
arrest of the process of development, by no means 
necessarily coexistent with diminished growth. '1 1" 
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, Substitu- 
tion, &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 pur- 
poses 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 cmite 
conceivable that opposition or superposition of organs 
may occur without the intervention of any such opera- 
tions. 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 alter- 
nate arrangement. Coalescence of the vascular 
bundles in an unusual manner, and an irregular dis- 


position of these cords have also been considered to 
bring abont 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, occa- 
sionally produce similar deviations, the nature of 
which is usually easily detected. 

Co-relation. — The importance of this subject first pro- 
minently brought into notice by Geoffroy St. Hilaire 
gains in force daily. Rarely is a malformation an iso- 
lated 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 recep- 
tacle. Let an ordinarily concave thalamus remain, from 
defective development, flat, and how great the change 
in the appearance of the flower. Let the usually con- 
tracted 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. 1 
Not only is the whole symmetry changed, but the 
physiological operations carried on in the flower un- 
dergo corresponding alterations. 

1 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 Avith perfectly green, long, tubular, erect, not 
horizontally spreading flowers. 


There are certain co-relations which do not appear 
to have hitherto attracted the attention they merit; 
sncli, for instance, is thai 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 in- 
definite 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 pro- 
longation 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 succes- 
sion, and, in consequence, arranged spirally. In the case 
of simultaneous development we meet with a repe- 
tition of whorls, as in what are termed hose-in-hose 
flowers (flores duplicati, triplicate, &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 wonld 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 

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 ex- 
pedient 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. 

0LA8STFICAT1 ■ \&9 

Take fche characters of exogens as distinct from 
end< even under ord 

lute distinctioi] can be drawn befcw m. Thereare 

plants normally of an intermediate character, whili . 
take exceptional instances, there are exogens with the 
Leaves and flow* adogens, and endogi out- 

ward or 3 them to exo- 

gens. Diclinous or monochlamydeous plants owe their 
imperfect conformation to sup] n, and may become 

structurally complete by a species of peloria. Struc- 
turally hermaphrodite flowers become unisexual 
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 recep- 
tacle that bears them, or by the arrested development 
of an ordinarily cup-like receptacle. 

How the relative position of the carpels and the 
calyx maybe 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. 1 Regular and irregular peloria, too, s< 
to show how slight are the boundai . t only 
between different genera, but also between different 

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 ? 

1 An illustration of this latter nature in the case of a cherry, which 
was surmounted by the calyx lob ly as in the eas..- of a prffha- 

is been given at p. 42 


Such questions cannot receive at present any satis- 
factory 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 denned 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 the 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. 



In ordinary language, the epithet double flowers is applied to fkn, 
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 vai'ious 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 

1 This appendix forms a portion of a paper published in the ' Pro- 
reedingsof the International Botanical Congress,' London, 18G6, p. 127, 
and which it has been deemed advisable to reproduce with sundry 
additions and modifications. 


obtain plants bearing double instead of single flowers. There is 
first the crossing of single flowers with double ones, effected by plant- 
ing 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 recog- 
nised 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 facility 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 1 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 green- 
houses, 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 sepa- 
rate those which would furnish double flowers from those which would 
produce single ones." 

' Traite des Giroflces,' per E. Chat* 


M. Chate's method, which he calls the French one, gives b1 ill 
results, viz. : 80 per cent, of double flowers, and these produced by very 
simple means. "When my seeds," he observes, "have been chosen 
wit it 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 tea 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 

Signor Biganionti, 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 occa- 

1 Leading Article in the ' Gardeners' Chronicle,' p. 74, 1866. 


sionally 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- 

" In the year 1820 we several times visited a garden in the neighbour- 
hood of Yienna, 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 conchtded that 
in the case of a plant imported from thence, and exposed to such 
different climatical influences, the origin of the greater or less imper- 
fection 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 


. transformed into d- - i Is it. therefore, remarkable that a plant 

intr m Japan int> Europe, expoe influences of this 

I diversity of climate, should produce imperfect sexual «.rgans 

incapable of further propagating the plant from Beeds p A rich soil, with 

the:. amount of moisture, will ] f double flowa 

Mr. Darwin - a peculiar form of Gen' reUa, in which 

the parte of the flower were more y compact . _ ■ 

tions of purple scales in great numbers. A similar condition is, indeed. 
not tmc«.>mm>.'n in this plant, and, as Mr. Darwin also remarked, on 
hard, dry. bare, chalky banks, thus bearing otit the views expressed by 
the writer in the ' Gartenzeitung ' just cited. Some double flow- 
PotentiUo £ und growing wild near York, and transmitted to the 

writer by s\ ondent, 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 ea 
sure to the sun, was particularly dry. 

On the other hand, the double-flowered Card . which is 

tonally found in a wild state, always grows in very wet pla 

Of late years a remarkable double-flowered race of P, 
has been obtained. In particular, Messrs. "Windebank and Kingsbury, 
of Southampton, ha .led in raising a set of plants in which the 

flowers a: uble and very attractive in a floi I view. 

The corollas in these flowers are not merely duplicated, but froru their 
inner surf ng, in some cases, funnel-shaped or tubular petals 

(p. 31-5), so regular in form as quite to resemble a perfect corolla. These 
tubes are attached to the inner side of th I 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 nume- 
rous ovules, hence it was at first surmised that these plants were 
obtained and perpetual the application of pollen from single 

flowers to these double- flowered varieties. 

The raisers of this fine race however assert that "the double 1: 
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 r 

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 

1 CM . tenzeitung.' 18 

- Gard. Chron.,' 1843, p. 628 

496 APPENDIX. • 

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. 1 

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. 

The 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 con- 
trasted 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 cultiva- 
tion, 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 impor- 
tant 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 first 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 2 Sometimes the flowers on a particular 
branch are double, while those on the rest of the plant are single. 3 
On these points, the evidence furnished by a double white hawthorn 
in the Royal Botanic Gardens at Edinburgh is important. Professor 

1 'Gard. Chron.,' 1887, p. 381. — Art. " Chinese primroses." 

2 See also p. 79, fig. 36. A similar flower is figured in ' Hort. Eystett. 
Ic. Arb. Vern.,' fol. 5. "Fructus nondum observatus est fortassis ali- 
mento uberius in flores refuso, nullus sperari possit." 

3 See De Candolle, « Plant. Rar, Genev.,' 1829, p. 91 ; and Alph. de 
Candolle. ' Geog. Bot.,' p. 1080. 


Balfour kindly wrote as follows in reply to an inquiry respecting tin's 
plant: — " A double white hawthorn in the Royal Botanic Gardens pro- 
duced 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 (lowering 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 pro- 

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 ob- 



jections raised by the present writer and others, and examines suinc 
of the alleged exceptions. Some of these the Belgian savant finds to 
prove his rule, inasmuch as although there is a co-existence of varie- 
gated 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 case3 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, Althaa, 
Pceonia, Rosa, Ranunculus, Serissa, Saponaria, etc., and probably the 
art of the cultivator would speedily be successful in raising other ex- 
amples, 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. 1 

Mr. Darwin 2 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 vege- 
tation ; 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 vena- 
tion, 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 pro- 
vided by cultivation, produce double flowers ? To this question, accord- 
ing to our theory, the reply would be that the quantity of food is 
excessive, more than the plant can properly digest ; and hence vegeta- 

1 See ' Gardeners' Chronicle,' 1868, p. 1113. 
- Ibid., 1813, p. 628. 

API . 

fcive 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 i with an excessive Btipply of 
food, which it cannot assimilate, is in many respects similar to that 
-ults 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 im- 
paired, or if the plant be pampered, it is no -wonderful thing that sterility 
shoidd 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 prolifi- 
cation, 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 develop- 
ment of leaves, in which all the parts — sheath, stalk, and blade — are 

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 Compositce, Viburnum Hydrangea, &c, are excluded. 


Clematis Viticella, Linn., S. Europe, 
florida, Thumb., Japan. 
Fortunei. Moore, Japan, 
patens, Dcsne. 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, X. America, Siberia. 

sylvestris, Linn., S. Europe, Siberia. 



Hepatica triloba, Chaix., Europe. 
Ranunculus bulbosus, Linn., Europe, N. Amer. 

repens, Linn., Europe, Siberia, N. Anier. 
acris, Linn., Europe, Siberia, 
aconitifolius, Linn., Europe. 

graniineus, Linn., Italy, France, Portugal, Switzerland, 
bullatus, Linn., S. Europe, 
asiaticus, Linn., The East. 
Ficaria ranunculoides, Mcench., Europe. 
Thalictrum anemoides, Michx., N. America. 
Caltha palustris, Linn., Europe, Asia, N. America. 
Trollius europseus, 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, Betz., Europe, 
tenuifolia, Linn., Tauria. 
albiflora. Pall., Siberia, 
paradoxa, Andr., S. Europe. 

Nelumbium speciosum, Willcl., Africa, Asia. 

Berberis, sp. cult. 


Papaver Rhoeas, Linn., Europe. 

bracteatum, Lindl., Russia. 

somniferum, Linn., S. Europe, Asia Minor, Egypt. 
Chelidonium majus, Linn., Europe, Asia. 
Sauguinaria canadensis, Linn., N. America. 
Podophyllum peltatum, Linn., N. America. 


Mathiola incana, 11. Br., Mediterranean. 

glabrata, D. C. 

annua, Siceet, 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, B. Br., Europe. 



Siaapie arvcnsis, Linn., Europe. 
Brassier oleracea, Linn., Europe. 


Helianthemum vulgare, Sjwch., Europe, N. Africa. 


Viola odorata, Linn., Europe, Siberia, 
grandinora, Linn., Europe, 
tricolor, Linn , Eui'ope. 


Dianthus barbatus, Linn., France, Germany, 
chinensis, D. C, Cliina. 
Poiretianus, Scringe, ? 
Caryophyllus, Linn., France, Italy, 
arboreus, Linn., Crete, 
bybridus (gardens). 
corymbosus, Sibth., Asia Minor, 
plumarius, Linn., Europe, Siberia, N. America, 
deltoides, Linn., Europe. 
Saponaria officinalis, Linn., Europe. 
Lychnis sylvestris, Sclikr., Europe, 
vespertina, Linn., Europe, 
flos cuculi, Linn., Europe. 
Viscaria, Linn., Europe, 
cbalcedonica, Linn., Japan, Asia Minor. 
Silene inflata, Sm. ; var. maritima, D. C, Europe. 

Sagina procumbens, Linn., Europe. 


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. 


iEsculus Hippocastanum, Linn., Europe, N. America. 


Geranium pratense, Linn., Europe, Sibei'ia. 

sylvaticum, Linn., Em'ope. 
Pelargonium zonale, Willd., S. Africa. 
Tropaxyhmi majus, Linn., Peru. 
minus, Linn., Peru. 



Oxalis cernna, Thumb., S. Africa. 
Impatiens Balsamina, Linn., E. Ind. 


Camellia reticulata, Lindl., China. 

Sasanqua, Thanh., China. 

japonica, Linn., Japan. 
Thea maliflora, Seem., Japan. 


Citrus Aurantium, Linn., Asia, South Europe. 


Trifoliuni repens, Linn., Europe, S. America. 
Medicago sp., ., Europe. 

Ulex europseus, Link., Europe. 
Spartianthus junceus, Linn., S. Europe. 
Clitoria Ternatea, Linn., E. India. 
Orobus viscoides, D. C, Croatia, &c. 

vermis, 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. 


Rosa lutea, Hill., Europe. 

cinnaniomea, Linn., Europe, N. America. 

spinosissima, Linn., Central Asia. 

Carolina, Linn., K". America. 

villosa, Linn., Europe, Central Asia. 

centifolia, liinn. 

damascena, Linn., Syria. 

rubiginosa, Linn., Europe, Asia, N. America. 

moschata, Ait., Madeira, N. Africa, 

canina, Linn., Europe. 

alba, Linn., Europe, Caucasus. 

indica, Linn., China. 

niyea, D. C, China. 

Eglanteria, Linn., Europe. 

gallica, Linn., Europe, Caucasus. 

pimpinellifolia, Linn., Europe, Central Asia, 

Banksiffi, J?.. Br., China. 

sulphurea, Ait., East. 
Rubus fruticosus, Linn., Europe. 

rosifolius, Linn., Mauritius, E. India. 
corylifolius, Smith, Europe. 
csesius, Linn., Europe. 
Kerria japonica, B. C, Japan. 


Spiraea Filipendula, Linn., Europe. 
Ulmaria, Iiinn., Europe. 

Srunifolia, Sieb. et Zucc, Japan. 
I,eevesii, Lindl., China, 
strobilacea, Sieb. et Zucc, Japan. 
Fragaria vesca, Linn., Europe, N. America. 
Potent ilia alpestris, Hall. /., Europe. 

reptans, Linn., Europe, Asia. 
Toruientilla, Schranh, Europe, Asia. 
anserina, Linn., Europe. 
Geum rivalc, Linn., Europe. 


Crataegus Oxyacantha, Linn., Europe. 

Crus galli, Linn., N. America. 
Cydonia japonica, Pers., Japan. 
Pyrus communis, Linn., Europe. 

Malus, Linn., Europe. 
Eriobotrya japonica, Lindl., Japan. 


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, TJiv.nb., China, Japan. 

iusititia, Linn., Europe. 

triloba, Lindl., China. 


Myrtus communis, Linn., S. Europe. 

Punica Granatum, Linn., S. Europe, Marocco. 


Philadelphus coronarius, Linn., S. Europe. 
Deutzia crenata, Sieb. et Zucc., Japan. 


Fuchsia globosa, Lindl. (and var. hort. pi.), Mexico. 
Epilobium tetragonum, D.C., Europe. 
Clarkia pulchella, Pursh., California, 
elegans, Douglas, N. America. 


Portulaca grandiflora, Hook, Chili. 

Ribes sanguineum. Pnrsli., N. America. 




Saarifraga granulata, Linn., Europe. 

Daucus Carota, Linn,. Europe. 


Ixora grandifiora, Be Cand., E. Iudia. 
Serissa fcetida, Comvi., China, Japan. 
Gardenia Fortuniana, Hook., China. 

florida, Linn., China, E. India. 

radicans. Tlmnb., Japan. 


Lonicera Periclymenum, Linn.. Europe. 
Sanibucus nigra, Linn., Europe. 


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. 


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. 

Epacris impressa, B. Br., Australia. 


Primula villosa, Jacq., Europe. 

Auricula. Linn,, Europe. 

denticulata, Smith, E. India. 

acaulis, Jacq., Europe. 

elatior, Jacq., Europe. 

pramitens, Kcr. = sinensis, Lin ell, China, 


Lyaunachia Nnrnmnlaria, Boem et 8chuU., Europe. 
Anagallis tenella, Linn., Europe. 


Jasminum officinale, Linn., S. Europe. 
Sambac., Ait, E. India, 
hirsuturn, Hook., China. 
grandiflorum, Lindl., S. Europe. 


Syringa persica, Linn., Persia. 

vulgaris, Linn., Europe, Persia. 


Vinca minor, Linn., Europe, 
major, Linn., Europe. 
Nerium odoi-um, Ait., E. India. 

Oleander, Linn., S. Europe. 
Tabernaemontana coronaria, Willi., E. India. 
Allamanda cathartics, Aubl, S. America. 


Calystegia sepium, i?. Br., Europe, America, Asia. 

pubescens, Lindl., China. 
Convolvulus tricolor, Linn.. S. Europe. 
Ipomcea pandurata, Meyer, S. America. 


Datura comigera, Hook., Pern. 

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., Em-ope. 

Gentiana Amarella, Linn., Europe. 

Orobanche sp. 



Minrulus luteus, Linn., Chili. 
Antirrhinum majus, Linn., S. Europe. 
Digitalis purpurea. Linn., Europe, 
Linaria vulgaris, Mill., Europe, N. America. 
Veronica, sp. 
Calceolaria, var. cult. 





Achimenes longiflora, D. C, Mexico. 
Gloxinia var. hort. 


Clerodendron fragrans, Willd., Japan. 
Verbena var. hort. 

Mirabilis Jalapa, Linn., Trop. America. 


Lauras nobilis, Linn., S. Europe. 

Sassafras, Linn., N. America. 


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 Ksernpferi, Siebold, Japan. 


Galantbus 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. 

Tazctta, Linn., S. Europe. 

poculiformis, Salisb., S. Europe. 


Orchis Morio, Linn., Europe. 

mascula, Linn., Europe. 

pyramidalis, Linn., Europe. 
Ophrys fucifera, Linn., Europe. 
See also pp. 380, 509. 


Hydrocharis Morsus ranse, Linn., Em-ope. 



Asphodelus luteus. Linn.. S. Europe, 


Tulipa Gesneriana, Linn.. Asia Minor. 

sylvestris, Linn., S. Europe. 
Seilla autumnalis, Lirm., Europe. 
nutans, Smith, S. Europe. 
Oonvallaria majalis, Linn., Europe, America. 

Polygonatum, Linn., Europe. 
Trillium grandifioruni, Spreng., America. 
Fritillaria Meleagris, Linn., Europe. 
imperialis, Linn., Persia. 
Lilium Martagon, Linn., Europe. 

candidum, Linn., Syria, Persia. 
Hyacinthus orientalis, Linn., East. 
Poliauthes tuberosa, Linn., E. India. 
Hemerocallis disticha, Don., Nepal. 
Kwanso, gardens, 
fulva, Linn., S. Europe. 


Colcliicum autumnale, Linn., Europe. 
Tofieldia calyculata, WaJil., Europe. 


Sagittaria latifolia, Willcl., N. America. 

sagittifolia, Linn., Europe, Asia, America. 



virginica, Linn., N. America. 





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 

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 Can- 
dolle, 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 specta- 
bilis shows two rows of carpels, placed one above another. The arrange- 
ment 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 par- 
tially coherent or sometimes quite distinct. The placentation is con- 
stantly 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 Yegetale," 
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 

NOTE. 509 

adventitious and imperfect flower-bud, as in the Phi mis, mentioned at 
p. L19. 

MoncBcioua Misleto, p. L93. — In this specimen, exhibited at one of the 

meetings of the Scientific Committ f the Royal Horticultural So< 

in L869, there were both male ami female dowers "n the same bush. 
plant was of the male sex, with numerous long slender whip-like, somewhat 
pendulous, branches bearing comparatively large broad yellowish l-'aves, 
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 ami 
immature female flowers. There was no evidence of grafting or para- 
sitism 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 
ricrasma, in which the leaves are impari-pinnate and spread horizon- 
tally, an adventitious leaflet was observed to project at right angles 
to the plane of the primary leaf. It emerged at a point nearly corre- 
sponding to that at which the normal pinna? 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, corres- 
ponding to the upper face of the normal leaflets, was directed towards 
the axis, i. e., away from the corresponding portion of the neighbouring 
pinna?, 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 
pinna?., 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 exag- 
gerated development of this gland, but no clue was afforded as to the 
origin of the ascidium. It was not practicable to examine the arrange- 
ment of the vascular bundles in the rachis. 

Additional labclla in Phaius. — A flower of Phaius cjrandiflorus was 
found in the same condition as the Catasetum, mentioned at pp. 291 and 

Tubular stern. — 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. Bollci, deserves 
notice from its beaiing on the question of such structures as the calyx- 
tubes, the hip of the rose and such like, see pp. 391, 482. In this plant 

510 NOTE. 

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 with- 
drawing it from the hand. 

The plant in question thus furnishes an actual illustration of the 
supposititious case mentioned at p. 4S2. 

Double flowers, see pp. 499, et seq. — The following species may be added 
to those already recorded : Lychnis coronaria, Hibisms mutabilis, Lotus 
major, Pisum sativum, Godetia sp., Ipomcea purpurea, Convolvulus tm 
TIeliotr opium peruvianum, THUivm grcmdiflorum, and Phaius grancli- 


Abortion, 455, 467 

of axile organs, 455 
cal v x . 

corolla, 460 
iiidusiuni, 467 
leaves, 458 
ovules, 466 
perianth, 460 
pistil, 46i 
receptacle, 457 
stamens, 463 
Acauleseence, 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 
meiotaxv 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) 

Autopbyllogeny, 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, 393 

Axile organs, see Axes 


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, pro- 

variations of, 336 
Bulbs, displacement of, 84 

multiplication of, 172, 350 
Burrs, 347, 420 


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), ad- 
ventitious, 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, 

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, 48 1 
inflorescence of, 245 
scales of, 192, 245, 484 
Compensation, 488 
Connective, petalody of, 293 
Consistence, alterations of, 432 
Contabesceuce, 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 


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, So- 
Displacement, 84 
of bulbs, 84 
carpels, 96 
flower-parts of, 91 
inflorescence, 85 
leaves, 87 
ovules, 96 
placentas, 96 
Distension, 419 


Ectlastesis, 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, 415 
sepals, 448 
stamens, 453 



Enlargement, 1)7 

of andrcecium, ISO 

axile org ins, 1 1 8 

buds, L20 

flower stalk, 1 u 1 

fruit, i::l 

gynoecium, 130 

perianth, 12 s . 

placenta, 125 
Epanody, - 
Epistrophy, 226 
Etiolation, ! 
Aversion, 204 
Excrescences, 1 It (see Enation) 


Fasciation, 1 1 

plants affected with, 20, a - 

Fastigiation, H' >2 
Ferns, crested, 63, 1 17 

depauperat -d. 

exindnsiate, 467 

supra-soriferous, 189 
Filaments, see Stamens 

petaloid, : 

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, 32S 
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, 10 1 

double, 490, 510 

hermaphrodite, 196 

homomorphic, 188 

increased number of, 390 

mutilated, 103 

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, 

Foliar organs, cohesion of, 21. 2", 

(■nation from, i !•"> 

fission of, 'il 

suppression of, 896 
Form, alterations of, 213 

juvenile, persistence of, 217 
Erondescence, 2 11, 27'.', Bee Phyllody, 

Fruit, adhesion of, 1 1 

enlargement of, 431 

Gemmae, formation of. 173 
Glands, formation of, 173 
Gnanrs, 10s, 347, U7, U9 
Grafting, 53, 56 
Greffe des Charlatans. 56 
Growth interrupted, 327 

irregular, 22S 
Gymnaxony, 211 
Gynantherus, 305 
Gyncecium (see pistils) 

enlargement of, 430 

meiophylly of, 399 

meiotaxy of, 4 16 

pleiotaxy of 

polyphylly. of, 363 

suppression of. 406 


Hairs, formation of, 472 

Hermaphroditism, 197 

Heterogamy, 190 

Heteromorpby, 311 

Heterophyllyi 330 

Heterotaxy, 166 

Homology, 47'j 

Homomorpliy. 188 

Hose in hose corollas, 291. 377 

Hypertrophy, 415, see Enlargement 

Independence, 58 

Indusium, abortion of, 467 

Inflorescence, displacement of, 84 
elongation of, 43 i 
prolification of, 102, 115 

Interrupted growth, 327 

Inversion of organs, 206 

Irregularity, 213 

Irregular growth, 228 


Knaurs, (see gnaurs) 
Kail, 426 




Laciniation, see fission 
Layering, 156 
Leaders, formation of, 203 
Leaf-slieath, 477 
Leaves, see foliar organs 

abortion of, 458 

adhesion of, 
by surfaces, 33 
to stem, 34 

adventitious, 162—165, 500 

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 


Meiotaxy of andrcecium, 405 
of calyx, 403 
corolla, 403 
gyncecium, 405 
Meiophylly of andrcecium, 398 
of calyx, 397 
corolla, 397 
gyncecium, 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, pleio- 

of bracts, 358, 371 
bulbs, 350 
cotyledons, 370 
embryos, 369 
florets, 351 
foliar organs, 352 
whorls, 371 


Xepaul Barley, 1 7 I 
Number, alterations of, 34 1 

increased, 343, 353 

diminished, 392 


Orchids, andrcecium of, 380 

prolification in, 153 
Organs, rudimentary, 469, 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 


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, 390 
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, 2S 
enation from, 448 
fission of, 66 
phyllody of, 251 
staminody of, 298 
tubular, 23, 314 
Phyllode, 328 



Phyllody, 240 (see Virescence, Ohio- 
of accessory organs, 272 
bracts, 242 
calyx, 2 1 l 

plants subject to, 25(1 
iii Conifers, 2 15 
of corolla, 25 1 

plants Bubject bo, 252 
of ovules, 262 

plants subject to, 271 
pistils, 256 

plants subject to, 261 
stamens, 2."):$ 

plants subject to, 256 
Phyllomania, 352 
Phyllomorphy, see Phyllody 
Phyllotaxy, 1, 320 
Pistil, abortion of, 46 I 
cohesion of, 29 
petalody of, 296 
Btaminody of, 298 
Pistillody, 302 

of ovules, 268, 310 
perianth, 302 
sepals, 302 
stamens, 303 

plants subject to, :;lo 
Pitchers, 30, 313,500, see Ascidia 
Placentation, changes in, 96, 508 
bibliography, 100 
nature of, 483 
Placenta, elongation of, 439 

enlargement of, 421 
Pleiomorphv, 228 
Pleiophylly, 353 

of androecium, 375 
bracts, 371 
calyx, 374 
corolla, 376 
gynoecium, 388 
perianth, 375 
Plica, 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 andrcecium, 361 
of calyx, 359 
corolla, 359 

plants subject to, 360 
of flower, 363 
gynoecium, 363 

Polyphylly, bibliography, 864 
plants subject tn, 36 1 
Position, changes of, 83 
relative, IS.") 

Prolification, 100 

axillary, L38 

foliar, I 11 

lloral, 1 12 

plants affected with, I is 
bibliography of, 15 I 

complicated, 151 

of embryo, 155 

of flower, 1 15 

coincident changes, 128 
median foliar, 116 
median floral, 1 19, 608 

plants affected with, 137 

of fruit, 13t 

inflorescence, 102 
bibliography, 115 

median foliar, 103 
median floral, 105 
lateral floral, 107 
lateral foliar, 106 


Receptacle, abortion of, 45 1 

elongation of, 1 1 6, I I I 

spiral torsion of. 325 
Reflexion, 209 
Regularity, 213 
Rhizotaxy, 1 
Rose Willow, 166, 168 
Roots, adventitious, 156 

elongation of, 434 
Rudimentary organs, 469 


Sajut Valery Apple, 135, 282, 304, 375, 

Savoys, 426 

Scales, formation of, 164, 448, 470 
Scape, leaves on, 163 
Seeds, abortion of, 407 

union of, 50 
Sepals, adhesion of, to petals, 3 I 

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 



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 Andrcecium 

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, 29S 

Stasiruorphy, 216 

Stem, see axes 

Stipules, increased number of, 357 
Strawberry, Plymouth, 275 

blind, 195 
Suppression, 393 

of andrcecium, 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 

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 As 

cidia, Solenaidv, Spurs) 
Tubular petals, 314 

stamens, 316 


Union, 8 

Unisexuality, 195 
Uovoli, 420 


Varieties, dwarf, 411 
Venation, 338 
Virescence, 338 
Viviparous plants, 106, 168 


Warts, 444 

Wheat-ear carnation, 371 


[In the following Index the names of the orders that arc incidentally men- 
tioned 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 fre- 
quency 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.] 


Abies Brnnoniana, 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-platnnus, 20, 371 
Aceranthus, 225, 226 
Achimenes, 106, 114, 296 

longiflora, 506 
Aconitum, 148, 231, 359, 360. 3'J'J, 404, 
407, 464 

Napellus, 39, 44, 238 
Acorus, 225 

Actsea spicata, 251, 252 
Adenium obesum, 417 
Adenorophinm luxuriant, 25 I 
Adonis, 202, 295 

autumnalis, 500 

vernalis, 500 

Adoxa, 367 

Aerides odoratuin, 398 

.Eschynanthus, 44, 297 

iEsculus Hippocastanum, 50, 66, 295, 

369, 438, 459, 501 
Affonsea, 364 
Af'zelia, 397 
Agaricus, xxiii, Si- 
Agave, 177, 432 

Americana, 33, 45 
Angelica, 365 
Agrimonia, 137, 406 
Agrostemma, 148 

Githago, 271 
Agrostis alba, 169 
Ailanthus glandulosa, 21 
Aira alpina, 169 

ctespitosa, 169 
Ajuga lva, 404 

pyramidalis, 20 

reptans, 2 13 
Alcea, 149, 297 
Alchemilla minima, 171 
Aldrovanda vesiculosa, 86 



Alisma, 115, 329 
natans, 107 
parnassifolia, 167 

Alismace^:, 115 
Allumanda, 296 

cathartiea, 288, 390, 505 
Allium, 106, 114, 170, 299, 360, 365, 

fragrans, 369 

vineale, 150 
Allium officinalis, 269, 271 
Almond, see Amygdalus. 
Alnus, 349 

fruticosa, 192 

imperialis, 459 

incana, 21 

glutinosa, 66, 244, 349 

laciniata, 65 
Alopecurus pratensis, 169 
A 1 si ne media, 67, 252, 404 
Alstromeria, 319, 326 
Althsea, 295 

rosea, 20, 501 
Alyssum, 137 

incanum, 252 

AMARYLLIDACEJ3, 115, 138, 150 

Amaryllis, 307, 310, 432 
Ambrina ambrosioides, 397 
Amentace^:, 114, 435 
Amorpha, 21, 397 

fruticosa, 23, 30, 243 
Amtgdale^:, 500 ; see Rosacea 
Amygdalus, 122, 137, 155, 295, 297 
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 

phcenicea, 141, 253, 271, 441, 461 
tenella, 505 
Webbiana, 281 
Anomodon alternatus, 49 
Ananassa, 350 
Auchusa, 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 (seelfepatica) 
coronaria, 107, 250, 256, 339, 499 
hortensis, 107, 250, 339, 499 
japouica, 161, L99 
nemorosa, 25n, 256, :;.i!>. 199 
palmala, 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 
Antbriscus, 442 

Anthurium Scherzerianum, 358 
Antbyllis, 295 

Antirrhinum majus, 20, 22, 24, 31, 44, 
57, 73, 82, 104, 114, 121, 13], 

225, 226, 227, 230, 233, 235, 
238, 253, 272, 296, 301, 315, 
316, 318, 363, 365, 378, 398, 

Apargia, 114 

autumnalis, 20 
Apium, 82, 113, 150 

graveolens, 66, 158 

Petroselinum, 370, 437. 
Apple, St. Valery, 135, 282, 304, 375, 

Aquilegia canadensis, 500 

Skinneri, 266, 271 

vulgaris, 24, 70, 127, 136, 220, 

226, 252, 257, 260, 261, 271, 
280, 286, 287, 288, 293, 295, 
374, 390, 500 

Arabis, 148, 508 

alpina, 397, 461, 463 

pumila, 170 

sagittata, 44 
Araucaria, 245 
Arbutus Unedo, 291, 292, 296, 377, 

Arcbidendron, 365 
Ardisia serrulata, 369 
Arenaria serpylli folia, 461 

tetraquetra, 398, 399, 464, 469 
Armeria, 114 
Aristolochia Clematitis, 38, 45, 314 

caudata, 231, 238 

sipbo, 23 
Armoracia rusticaua, 64, 299 
Arnoseris, 114 
Artabotrys, 456, 508 
Artemisia, 405 
Artocarpus, 407 
Arum, 329 

maculatum, 66, 225, 227, 215, 35S 
Asparagus officinalis, 12, 19, 21, 320, 

325, 171 
Asphodelus, 138, 296 




Aspli. uli-lus lutens, 506 
ramosns, 298, 310 
Aspleniam Tricbomaoes, 190 

Astrantia, 1 ! I 

major, 868 
Athamanta, 82, 187, 1 10 

Cervaria, 250, 112 
Atriplex, 227 

hortensis, 221 
Atropa Belladonna, 1 1 
Aucnba, 21 

japonica, 21, 197 
Auuantiace.e, 137, 119, 502 
Avena, 319, 351, 391 

chinensis, 298 
Azalea, 35, 114 

glanca, 50 1 

indica, 11, 73, 166, 200, 289, 296 

nudiflora, 501 


Babingtonia, 185 
Bsackea diosuaifolia, 183 

1>\i.>aaii>'ejl:, 359, 104, 501 
Balsam, see Impatiens 
Bamboo, see Bambusa 
Bambusa, 307, 310, 324, 365 
Barbarea, 205 

vulgaris, 310, 500 
Barkhausia, taraxacifolia, 20 
Barley, Nepaul, 174 
Baubinia, 328 
1!egoxiace.e, 114 

Begonia, 31, 81, 100, 114, 162, 170, 

frigida, 199, 303, 310 

fuchsioides, 281 

phyllomaniaca, 170 
Bellevalia, 408, 461, 467 

comosa, 348 
liellis 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 


Bignonia, 272, 296, 327 
Bikkhia, 80 
Blitum, 45 

polymorpbum, 397, 458 
Bocconia, 224 

cordata, 310 

Bowiea rolnbilis, 409 

HoitAt.iN \< i: i ■:, l:<2, 138 

Boncbea hyderabadensis, 166 
Bougainvillea, 330 
Bracbytbeciam plumosnm, 40 
Brassica, 20, 136, 139, 205, 364, 367 

Napus, 27, 205 

oleracea, 30, 33, 66, 67, 106, 1 13, 
136, 141, 142, 1 is, 250, 252, 
261, 271, 280, 351, 408, 421, 
426, 112, 115,501,508 

Rapa, 181 
Breynia, 198 
Bromelia, 103 
Bromus velutinus, 358 
Bronssonettia papyrifera, 331, 459 
Bruniacese, 80, 81 
Brunia microphylla, 81 
Bryonia, 360, 367 
Bryopbyllum calycinum, 15S, 171, 183 

proliferum, 166 
Bryum csespititiuin, 49 
Bunias, 97, 136, 280, 360 
Bonium creticum, 159 

flexuosum, 20 
Bupleuruin, 140, 450 

falcatum, 15, 20 
Bctoaiace.e, 507 
Buxbaumia indusiata, 49 
Byrsoniraa, 137 



Cabomba aquatica, 458 
Cactace^e, 81, 113, 140, 39a 
Cactus, 160 
Cachrys taurica, 197 

Cajsalpinia, 365 

digyna, 43 
Cakile maritima, 246, 250 
Calantbe, 227, 398, 402, 5( is 

vestita, 30, 45, 227, 402, 508 
Calceolaria, 41, 11, 230, 233, 284, 296, 
397, 405, 406, 505 

crenati folia, 238 

floribunda, 316 

rugosa, 238 
Calendula, 114, 138, 339, 370 

officinalis, 252, 280, 339 
Calla palustris, 357 
Callitriche, 196 

autumnalis, 406 

vernalis, 406 
Caltba, 136, 148, 295 

palustris, 141, 250, 442, 5uo 
Call una, 296, 504 
Calycophyllum, 249. 283, 429 
Calvstegia, 114, 296 



Calystegia Sepium, 505 

pubescens, 505 
Camellia japonica, 288, 295, 297, 491, 
494, 502 

reticulata, 502 

Sasanqua, 502 
CAMPANULACE-aS, 80, 114, 127, 131, 

138, 139, 150, 334 
Campanula, 71, 72, 73, 82, 138, 150, 
250, 285, 296, 3G5, 367, 404, 
442, 472 

canescens, 403 

colorata, 403 

gloraerata, 242, 300, 504 

latifolia, 504 

Medium, 20, 37, 44, 61, 251, 448, 

persicifolia, 44, 284, 300, 429, 504 

pyramidalis, 281, 504 

Iinpunculus, 429 

rapunculoides, 20, 252, 300, 310, 

rhomboidea, 50 1 

rotunditblia, 377, 378, 504 

Tenorei, 504 

thyrsoidea, 20 

Tracbelium, 504 

Vidallii, 504 
Campanumaea, 80, 81 
Camphorosma monspeliaca, 456 
Canna, 285 
Cannabis, 82, 194, 197 

sativa, 81 
Cannamois virgata, 197 
Capparidace^:, 148, 390 
Caprifoliace^:, 45 
Capsella bnrsa pastoris, 298, 361 
Cardamine, 295, 357 

birsuta, 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 

csespitosa, 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 

Kulbocastanum, 159 

Carui, 244, 271, 285, 339, 437 
Caryophyllace^:, 99, 113, 120, 137, 
139, 140, 148, 250, 379, 397, 
398, 404, 406, 407, 410, 418, 
442, 443, 448 
Casuarina vigida, 325 
Cassia, 364, 369 

marylandica, 30 
Castanea vesca, 11, 66, 104, 114, 319, 

Catabrosa aquatica, 351 
Catalpa, 399 
Catasetum, 291, 296, 334 

eburneum, 384 
Cattleya ametbystina, 401 

Forbesii, 34, 384, 398 

marginata, 223, 2^7 

Mossise, 224, 227, 238 

violacea, 383, 397 
Caucalis leptopbylla, 33 
Caulophyllum. 75, 125 
Caylussa, 137 
Cedrus Libani, 61 


Celastrus, 149 
Celosia, 19, 20 
Centaurea, 37, 114 

collina, 34 

Jacea, 43, 243 

Scabiosa, 20 

moschata, 44 
Centranthus, 247 

macrosipbon, 250 

ruber, 42, 44 
Cepbalotus 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, 

avium, 262 

caproniana, 364 

Lauro-cerasus, 64, 66, 370 

vulgaris, 252, 262 
Ceratonia Siliqua, 30 
Cercis, 364 

siliquastrum, 325 
Chama?rops buinilis, 300 
Chaerophyllum, 113 
Cbeiranthus, 121, 131, 136, 148, 295 

Cbeiri, 20, 33, 35, 36, 250, 252, 
271, 404, 427, 500 

var. gynantberus, 305, 310 


■ r )21 

( Iheiranthus incanus, 250 

barbata, 238 
Chelidonium maj us, 66, 170, 280, 295, 

Ch BKOPOD] w i' I . 897, 405, 40G 
podium, 45, 365, 367 

glaucum, 397 

murale, 428 

Quinoa, 62, 66 

Vulvaria, 458 
Cbirita sinensis, 1 70 
Chloropbytum Sternbergianum, 107 
Cborozema ilicifolium, 21 
Chrysanthemum, 1(5, 72, 365 

indicnm, 20, 188, 172 

Leucantbemum, 20 
Clirysosplenium, 367 
Cicborium Intybus, 20, 44 
Cicata virosa, 406 
('inoiionacejE, 429 
Cionidium Moorei, 190 
Cirsium, 114, 138 

arvense, 250, 457 

lanceolatum, 20 

tricephalodes, 252, 339 
Cissus, 211 

ClSTACEiE, 137 

Cist us vaginatus, 473 
Citrus, 137, 149, 364, 453 

Aurantimn, 33, 35, 44, 56, 75, 
134, 303, 310, 335, 369, 388, 
389, 391, 502 
Clarkia, 24, 295 

elegans, 177, 503 

pulchella, 503 
Cleistauthus polystachyus, 198 
Clematis, 136, 148, 288, 295, 367 

fiorida, 499 

Fortunei, 499 

patens, 499 

Viticella, 28, 499 
Cleouie, 137, 148 
Cleonia lusitanica, 238 
Clerodendrou fragrans, 506 
Cliffortia, 396 
Clinacium dendroides, 49 
Clitorea Ternatea, 231, 238, 295, 502 
Clusia rosea, 11 
Cluytia semperflorens, 198 
Cneoruui, 365 
Cnicus palustris, 20 
Cnidium, 113 

Coba>a scandens, 73, 82, 272, 326, 365 
Coccoloba platycladon, 328 
Cochlearia Amnoracia, 64, 299, 310, 331 
Cocos, 115, 365 

nucifera, 429 





( '>'<li;rmii variegatum, 31, 314, 326, 

Coelebogyne ilicifolia, 369 

Cola acuminata, :s7o 

Colcbicnm antumnale, 45, 67, 73, 250, 

287, 296, 406, 407, 507 
Coleus,365, 159 
( lolumnea Scbiedeana, 226 
Columbine, see Aquilegia 
Colutea, 165 

Commelyna, 73, 296, 507 \ci:j;, 215, 507 
Composite, 72, 73, 86, 107, 114, 
131, 138, 165, 223, 226, 
339, 406, 407, 430, 434, 
439, 4 12 
Conceveiba macrophylla, 198 
Conifers, 56, 65, 103, 114, 191, 

369, 435 
Conium maculatum, 114 
Conostepbiura, 120 
Convallaria maialis, 73, 150, 250, 
360, 367, 375, 400, 442, 

Polygonatum, 507 
Convoltulacej:, 114, 137, 150 
Convolvulus, 73, 114, 137, 142, 
296, 510 

arvensis, 20 

Sepium, 20, 108, 250 

tricolor, 505 
Conyza squarrosa, 20 
Corallorbiza innata, 238 
Circeia, 410 

Coreborus acutangulus, 397 
Coreopsis, 20, 73, ll4, 138 

Druramoudi, 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, 213 

tuberosa, 235, 336, 238 
Corylus Avellana, 21, 31, 48, 66, 114, 

349, 354, 361, 365, 368 
Cotoneaster micropbylla, 21 
Cotula fcetida, 19, 20 
Ceassulace^:, 113 
Crassula, 113 

arborescens, 26, 31 
Crataegus, 82, 113, 149, 295, 364, 404, 

Crus galli, 503 

monogyna, 44, 400 



Crataegus Oxyacantba, 57, 66, 78, 317, 
370, 503, 508 
tanacetifolia, 107, 162 
Crepis, 271 

virens, 20 
Criuum, 432 

Crocus, 29, 35, 45, 67, 287, 289, 296, 
319, 361, 399, 400, 434, 462 

aureus, 506 

nudiflorus, 302 

pusillus, 506 

vermis, 506 
Crozophora tinctoria, 198 
CKtciFERJE, 73, 76, 98, 113, 136, 139, 
141, 148, 257, 297, 364, 379, 
406, 410, 428, 442, 500 
Crvptomeria japonica, 103, 114, 245, 

Cucubalus, 149 
Cucumis, 36, 82, 138, 248, 259, 326, 

CrcrEBiTACEiE, 71, SO, 81, L37, 217 
Cucnrbita, 197, 201, 250, 3u7, 310, 365, 

Cuphea miniata, 211, 424 
Cupressus funebris, 218 
Ctcade.e, 56, 369 

Cyclamen, 67, 104, 114, 296, 319, 359, 

1 in eari foli ura, 329 
Cyclodon, 80 
Cydonia vulgaris, 71, 79. -'Jo, 123 

japonica, 5l *3 
Cynancbum fuscatuin, 369 

nigrum, 369 
Cynosurus cristatus, 169 
Ctpeeacex, 115, 138. 15m, 169, 350 
Cypripedium, 27, 92, 381, 386 

candidum, 401 

Hookera?, 386 

insigne, 91 
Cyrianthus, 177 
Cytisus, 113, 295, 336 


Laburnum, 21,66, 157,189, 222, 
220, 231, 238, 356, 159 

nigricans, 15, 101 


Dactylis, 1 1 '< 

glomerata, 169 
Dahlia, I '. 72, 1 J8, 2)1, 133 

variabilis, 20 
Danae, 0<j 
Daphne indica, 21 

odora, 21 
Datura, 285, 201, 296, 3 

Datura arborea, 505 
cornigera, 505 
cblorantba, 505 
fastuosa, 296, 379, 450, 505 
hurailis, 505 
Daucus Carota, 53, 57, 82, 113, 

124, 125, 137, 149, 2 11, 

252, 256, 260, 262, 296, 

339, 365, 368, 369, 370, 

Delphinium, 44, 136, 148, 225, 

286, 295, 344, 364, 367, 

388, 399, 407 
Ajacis, 250, 252,261,271, 339 
amcenum, 261 
cheilanthum, 500 
Consolida, 373, 500 
crassicaule, 252, 256, 261, 27L 
dictyocarpum, 271, 432 
elatum, 20, 126, 237, 238, 

267, 271, 309, 310, 339 
elegans, 500 
grandiliorum, 500 
peregrinuin, 219, 226 
Dendrobium, 227 
nobile, 94, 398 
normale, 22 1, 383 
Desmodium canadense, 271 

marylandicum, 467 
Deutzia, 295 

crenata, 503 
Dianthus, 67, 113,121, 129, L37, 

145, 146, 119, 1.66. 261, 

289, 295, 207. 310, 360, 

371,379, 307, 171 
arboreus, 501 
barbatns, 325, 404, 501 
Caryophyllus, 501 
corymbosus, 501 
deltoides, 501 
hybridus, 501 
plumarius, 501 
Poiretianus, 501 
sineusis, 370, 501 
Dictamnus, 121, 122, 137, L40 
albus, 256, 271 

Fraxinella, 252, 262. 271. 278 
Dielytra, 236, 237 

Lis lutea, 60 
orieutalis, 238 
purpurea. 20, 40, I !, 73, 98, 

129, 137, 150, 220, 233, 

296, 29S, 315, 365, 373, 

159, 4.72, 505 



Muscipula, 310 
Diosina, 300 
Dipbaca, 365 















Diplotaxis, L86, l is, B64 

moralU, ~'>~, 158 

tenuifolia, 78, 860, 261, 271, 280, 
397, 398, 180 
[)IP8ACA< I B, B6, H '7, 114, 133 
Dipsacua, U9, l-'.' 

fallonnm, 20, 62, GO, 281,»820, 
321. 325 

Qmelini, 325 

pilosus, 20, 325 

sylvestris, 1<», 20 
Dipterocarpus, 2 19 
Ditaxis lancifolia, 380 
Dodecatheon, 133 
Dodonsea riscosa, 20 
Drul .a, 364 

Dracocephalum austriacum, 238 

mold ivicum, 20 

Bp < iosum, 320, 325 
Dracontium pertuBum, 25 
Drosera intermedia, 170, 265, 271, 277, 


Ebenus cretica, 26 

Eccremocarpus scabcr, 222, 226, 326 
Ecbeveria, 113 
Echinophora maritima, 252 
Echium pyrenaieum, 20 

simplex, 20 

vulgare, 374 
Echinocactus, 149, 178, 417 
Blegia, 115 

Euipetrmn nigrum, 197 
Encamptodon perichsBtialis, 17 i 
Epaceidace.e, 120, 137 
Epacris, 103, 137 

impressa, 61, 379, 501 
Epidendrum, 114 

elongatum, 107 

Stamfordianum, 401 
Epimedium, 23, 226 

Musschianum, 390 
Epiphyllum, 323 
Epipactis palustris, 325 
Epilobium, 81, 137, 273 

angnstifolium, 20 

hirsutum, 24G, 250, 252, 256, 262, 

palustre, 271 

tetragonum, 503 
Epi8cia bicolor, 170 
Equisetacej:, 189, 350 
Equisetnm, 61, 325 

rluviatilc, 325 

liniOBum, 325 

Telmateia, 323, 325 

Eranthis hyemalis, 23, 70 
BBICAO&fi, I l I, L19, L87 
Erica, 21,73, 187, 296 

cinerea, 50 1 

hyemalis, 378, 50 l 

multiflora, 372 

Tetralix, 286, 810, 105, 406, 604 
Eriobotrya japonica, 2'j5 
Erodinm, 163 
Brvum Lens, 20, 25, 66 
Erucago, 136 
Eracaatrnm Pollicbii, 271 

eaiiariense, 280 
Eryngium, 113, 368, 412 

viviparnm, 104 
Erysimum, 136, 1 18 

Barbarea, 252 

cheiranthoidcs, 252 

officinale, 252 
Erythrochiton hypophyllanthus, 32, 

Esclinlzia crocea, 250 
Eucomis, 103 
Eugenia Jambos, 369 
Euonymna japonicus, 20 

latifolius, 369 
ErPHORBiACE.E, 114, 150, 369, 395 
Euphorbia, 114, 150, 365, 371, 395 

Characias, 20 

Cyparissias, 20, 2 1 1 

Esula, 198, 307, 310 

exigua, 20 

geniculata, 253, 256 

helioscopia, 56 

Lathyris, 2 i 1 

rosea, 369 

palustris, 106, 299 

Peplus, 162 

pusilla, 244 

sogetalis, 281 


Faba, see Ficia 

vulgaris, 397 
Fabiana, 237 
Fagus silvatica, 65, 66, li>7, 31c, 370, 

Festuca, 115 

nemcralis, 169 

ovina, 169 
Ficaria, (see Ranunculus Ficaria) 

rannuculoides, 70, 295, 500 

ElCOIOE.K, -13 

Ficus Carica, I 1 1. 201, 435 

stipulata, 332 
Filago, 1 1 1 

germanica, L08 



Filices, 21, 190, 447 
Fourcroya, 115 

Fragaria, xxxvi adnot., 250, 295 
alpina, 271 
botryforinis, 47 
monophvlla, 396 

vesca, 20, 70, 116, 195, 275, 281, 
406, 503 
Fraxinus excelsior, 13, 21, 66, 325, 396, 
Ornus, 21 
Fritillaria iniperialis, 21, 45, 296, 462, 
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, 442 
globosa, 503 
Funckia, 369 


Gagea, 365, 367 

arvensis, 375, SOS 
Gaillardia, 269, 271 
Galaothns, 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 
( raudichaudiese, 334 
Gardenia, 296, 377 

florid a, 504 

Fortuniana, 504 

radicans, 501 
Gaura biennis, 20 
Genista, 295 

tinctoria, 502 

sibirica, 502 

Scoparia, 502 
Gentianaceje, 137, 150, 505 
Gentiana, 71,73, 137, 150, 252, 296, 

Amarella, 166, 305, 310, 371, 505 

campestris, 250, 299, 412 

purpurea, 389 
Gebaniace-e, 113, 137, 501 
Geranium, 20, 137, 221, 246, 250, 252, 
292, 295, 418 

eolumbinum, 461 

nodosum, 34, 65 

pratense, 501 

svlvaticum, 501 

Gesnebace^e, 38, 114, 222, 505 
Gesnera, 33, 44, 95, 171, 339, 357, 427 

Geroldtiana, 88, 89 

zebrina, 355 
Geum, 121, 137, 465 

coccineum, 275 

fivale, 122, 130, 131, 250, 252, 281, 
Gilia capitata, 281 

glomeruliflora, 253, 271 
Gladiolus, 21, 296 

tristis, 506 
Glaucium luteum, 66 
Gleditscbia, 30, 177, 364 

triacanthos, 44, 48 
Glocbidion, 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 
Geaminacejj, 115, 138, 278, 350, 391 
Gratiola, 296 
Guarea, 508 
Gypsophila, 149 


Habenaria, 138, 238 

cblorantha, 382 
Halenia, 222 

heterautba, 222, 226 
Hedera Helix, 65 
Hedypnois, 114 
Heliauthemum, 132, 137, 295. 404 

vulgare, 501 
Helianthus, 38, 44, 66 

annuus, 20 

tuberosus, 20 
Hellebore, 23. 288 

fcetidns, 112 

olympicus, 284 
Heliotropium peruvianum, 510 
Helwingia, 174 
Heineroeallis, 138, 296, 310, 5U7 

disticba, 507 

fulva, 507 
Hepatica, 295, 463 

triloba, 500 
Heracleum, 82, 113, 137, 150,262, 365. 

Spbondylium, ^.'»2, 2r>6, 339^ 
Hermesia castaneifolia, 1'.)! 
Herreria parviflora, 141, 150 



ris, 19, 136, 295 

matronalis, 20, 252, 280, 825, 500 
Beterocentrou, :!•"> I 
HibifloUB, L87, 293, 295, 297, 3(30,510 

albus, 501 

flaveacens, 501 

llosn sinensis, 501 

Syriacus, 20, 501 
Hieracium, 138 

anrenm, 17 

proaltnm, 339 

l'iloM'lll, 20 

ambellatum, 20 

HirrOCASTANE.E, 501 

Hipppeaatrum, 296 

equestre, 506 
Hippuris, 196 

vulgaris, 325, 406 
Hodgsonia, 32<i, 17 J 
Holcus mollis, 169 
Honckenya peploides, 196, 106, 461 
Hordeum, 115, 351 

nepalense, 174, 175 

trifurcatum, 17 1, 175 
Huraulus Lupulus, 193, 244, 435, 472 
Hyacintlras, 138, 150, 262, 296, 360, 
361, 486 adnot 

comosus, 409 

orientalis, 21, 15, 48, 54, 172, 189, 
286, 299, 348, 507 

Pouzolzii, 170 
Hydrangea, 417 
Hydrocera triflora, 359 
Htdeocharidacej:, 506 
Hydroeharis, 296 

morsus ran®, 506 
Hydrocotyle, 113 


Hydrophyllum, 138 
Hymenocallis, 404 

aniericana, 462, 463 
Hyoscyamus, 430 
Hypericum, 442 

perforatum, 369, 458 
Hypnum triquetrum, 49 
Hypocliffiris, 73, 338 

radicata, 250, 339, 437, 457 
Hyssopus officinalis, 20, 325 


Iberis, 295, 364 

amara, 500 

umbellata, 500 
Ilex Aquifolium, 21, 66, 447 
Impatiens, 161, 231, 295, 299 

Balsamina, 238, 502 
Imperatoria, 82 

[ndigofera, 169 

Inula, 20 

[onopsidium acanle, o<'>- 
Ipomoea, 296, 510 
pandurata, 505 

Iimhac i:.k, 138, 506 
Irina, 6 I 

glabra, 65 
Iris, 138, 286, 296, 359, 360, 361, 305, 
401, 430 

K;i mpferi, 506 

sibirica, 506 

versicolor, 45 
Isocbilus, 386 
Ixia carminosa, 84 
Ixora, 296 

grandiflora, 50 4 

Jasione, 20, 114 
Jasminace^:, 137, 504 
Jasminum, 137, 296, 360, 400 

grandirlorum, 288, 505 

hirsutum, 505 

nudiflorum, 21 

officinale, 21, 505 

Sambac, 505 
Jatropha Pohliana, 254, 256 
Juglans, 244, 400 

nigra, 396 

regia, 66, 193 
JttncacejE, 115, 167, 169 
Juncus, 115, 317 

articulatus, 431 

conglomerates, 325 

uliginosus, 107 
Juniperus virginiana, 194 

sinensis, 217 
Jussieua, 81, 180 
Justicia oxyphylla, 25 


Kerria, 295 

Knautia arvensis, 20, 114 

Koehia Scoparia, 430 


Labiate, 138, 429 

Laburnum (see Cytisus), 65, 157, 189, 

222, 226 
Lacistema, 359 
Lactuca, 114 

sativa, 11, 20, 33, 44, 313 



Lauibevtia, 365 
Lampsana, 1 11 
Lamium, 73, 238, 361 

album, 62, 63, 66, 86, 409 

amplexicaule, 4< 1 1 

purpureuui, 66, 325,404 
Larix, 114 

europaa, 21, 90, 245, 435 

microcarpa, 192 
Latliyrus latifolius, 262 

tuberosus, 30 
Laurus, 296, 362 

uobilis, 506 

Sassafras, 250, 331, 506 
Lavatera trimestris, 20 
Lebeckea, 459 
Lecytbis, 149, 180 

Leguminosje, 48, 71, 73, 106, 113. 122, 
137, 139, 146, 147, 149, 272, 
276,429, 434, 11! 
Leitnera floridana, 194 
Leoutice, 125 

Leontodon, 17, 20, 44, 163, 243, 442 
Lepidinm, 148, 364 

sativum, 57 
Lepyrodia hermapbrodita, 197 
Leskea sericea, 49 
Leueantbemum, 86 
Leucobryum giganteum, 194 
Leucoium, 150, 296 

sativum, 84, 138, 350 

vernum, 350, 506 
Lilium, 73, 106, 115, 296, 367, 375, 

auratum, 73, 289, 400 

bulbiferum, 45 

candidum, 21, 286, 325, 375, 507 

cruentum, 21 

lancifolium, 35, 100, 113 

longirlorum, 310 

Martagon, 21, 286, 325, 507 

tigrinum, 306, 310 
Linaria, 137. 220, 230, 233, 296, 316, 
361, 365, 367, 405 

reruginea, 238 

cbalepensis, 238 

Cymbalaria, 238 

decumbens, 238 

Elatine, 238 

origanifolia, 238 

Pelisseriana, 238 

pilosa, 238 

purpurea, 20, 14, 238 

spuria, 238 

tripbylla, 238 

vulgaris, 162, 226, 234, 235, 238 
316, 374, 505 

triornitbopbora. 238 
Linum, 335 

! Linum usitatissimum, 20 
Liquidambar, 362 
Listera ovata, 308 
, LOBELIACE.E, 72, 11 1 
i Lobelia, 211, 424 

Lolium, 86, 113, 115, 351 

perenne, 61, 169, 325 
Lonicera, 38, 44, 73, 82, 226, 271, 281 , 
296, 297 

bracbypoda, 358 

Caprifolium, 408 

Periclymeuum, 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, 5' ):i 

uligiuosus, 363 
Lovvea, 396 
Lunaria, 136, 364 
Lupiuus, 106, 165, 226, 280 

polypbyllus, 238 
Lycaste Skinneri, 93, 95, 291 
Lychnis, 113; 137, 148, 295 

chalcedonica, 501 

Coronaria, 107, 510 

dioica, 67, 252, 262, 280, 401, Mil 

flos cuculi, 501 

sylvestris, 252, 501 

vespertina, 501 

Viscaria, 501 
I Lycium, 365, 367 

europasum, 250 
Lyeopersicum, see Solan inn 

esculentum, 389 
Lysimachia, 119 

Ephemerum, 271, 281 

nummularia, 505 

vulgaris, 20, 87 
Lythrum, 335 

Salicaria, 374 


Madenia, 403 
Majsa, 145, 371 
Magxoltacejs, 122 
Magnolia, 288, 388, 440 

Campbelli, 127 

fuscata, 304, 310, 427 
Maipighiace.e, 137, 331, 403, 401. 106 
Malus, 78, 79, 388, 389, see Pt/rus 
Maevace^:, 137, 149, 288, 292, 2'.»:., 

362, 395 
Malva, 295 

crispa, 448 

moscbata, 501 

rotuudit'olia, 501 

sylvestris, 252 

i\i.| \ OF s VMES OF PLANTS. 

■ r_L< 

ifera, L59, 369 
Marchantia, 17 I 
raavia, 23 
orabellata, :<::2 
MasdevalUa, 27, 95 
Mathiola, 186, 1 is, 295 
annua, 86 I 

incana, 20, 38, l i, 68, 299, 500 
Matricaria Parthenium, 2s] 
Maxillaria, 883 

Medicago, 137, 1 19, 262, 295, 864, 502 
topulina, 404, 132 
maculata, 218,271 
Bffelaato ua, 7 I, !-•">, I- I 
BCelia Azederacb, 21 
Meliantbus major, 298, 107 
Melilotua, 137, 139, 147,1 W, 262 
arvenais, 37 I 
leucantba, 132 
macrorbiza, 271 
officinalis, 40 I 
MelittiB, 361 
Mentha, 238, 361, 159 
aquatica, 325 
vividis, 325 
Mercarialis, 62, 19 1, 365 
annua, 19S 
perennis, 6G 
Blesembryanthemum, 26 
Metrosideros, 103 
Miconia, 355 
Mimosa, 365 

Lopbaiitba, 31 
Mimulns, 73, 284, 296 

luteus, 505 
Mnium serratum,49 
Mirabilis, 296,418 

Jalapa, 506 
Modecca, 326 
Mollngo Cerviana, 398 
Momordica Elaterium, 20 
Monarda fiatolosa, 298 
Moms, 193, 354, 459 
Mozinna, 399 

peltata, 194 
Musa, 245, 407 
Muscari, 408, 461 

eoniosum, 340, 467 
Mrsci, 17 ! 

Mussamda, 249, 283, 429 
Myosotis cKspitosa. 281 
palnstris, 375 
scorpioides, 20 
Myosurus, 440 
Myristica moschata, 194 
M yktaoeje, 80, 149, 362, 395 
Myrtus, 295, 310 
communis, 503 


Narcissus, 21, 67, 188, 286, 296, 860, 
365, 143 
aureus, 506 
biflorus, 84, 15, 506 
cbrysantbuB, 88, 15 
concolor, 506 
Cypri, 506 
Jonquilln, -V 16 

incomparabilis, 38, 15, 301, 506 
italicus, 506 
lobularis, 506 
major, 1 5 1 
montanu8, 301 
poculiforinis, 506 
poeticue, 24, 33, 301, 506 
pseudo-narcissus, 506 
Tazetta, 45, 300, 506 
Telamouius, 506 
Nasturtium, 136, 271 
ampbibium, 181 
Nelumbium, 295 

speciosum, 500 
Nepeta diffusa, 238 
Nepenthes, 328, 173 
Nephrodium molle, I 17 
Nerium Oleander, 62, C>G, 296. 301 , :i77 

odorum, 505 
Neuropeltis, 32 
Nicandra pbysaloides, 158 
Nicotiana, 23, 31, 73 

rustica, 281 
Nigellu, 220, 262, 295, 297, 364. 374, 
damascene, 226, 269, 271, 286. 375, 
Nuytsia, 371 
Ntmph.t-.ace^, 148 
Nympbaja, 50, 141, 143, 118, 285, 295 
alba, 162 
dentata, 256, 261 
guineensis, 170 
Lotus, 277. 280 


Octadenia, 364 

Odontoglossum Alexandra?, 387. 403 

Uro Skinneri, 463 
OEnantbe, 82, 114, 360, 365. 467 

crocata, 80, 359, 157 
(Enothera, 44, 367 

striata, 252 
Olea europcea, 21, 157, 420 
ONAGRACEiE, 81, 137, 406, 503 
Oncidium, 67, 114, 227 

ahortivum, I<!2 

hi color, 45 

Ceholleta, 107 



Oncidium cucullatum, 91 

heteranthuui, 224 

ornithorhyncum, 45 
Ononis minutissima, 404 

toonophylla, 398 
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 

Hionacantha, 178 

Salmiana, 107, 178, 179 
OECHiDACEiE, 27, 34, 42, 91, 112, 13 4, 
128, 138, 150, 153, 209, 223, 
290, 345, 360, 366, 367, 380— 
387, 397 
Orchis, 138, 150, 153, 154, 227, 296 

conopsea, 238 

latitblia, 238 

mascula, 153, 154, 238, 299, 387, 

rnilitaris, 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 
Oeobanchacea, 137 
Orobanche, 73, 137, 209, 296, 505 

gracilis, 442 
Orobus, 295 

vernus, 502 

viscosus, 502 
Ouvirandra, 458 


Oxalis, 295, 329, 404, 459 
Acetosella, 404 
carnea, 502 

Pteonia, 295, 300, 364, 399, 407 

Moutan, 500 

officinalis, 261 

paradoxa, 500 

tenuiflora, 500 
Palmacea, 155 
Panax, 331 
Papaver, 289, 295, 297, 473 

bracteatum, 28, 304, 310, 500 

nudicaule, 310 

orientale, 250 

Papaver Rhceas, 429, 458, 500 

somniferum, 28, 305, 310, 500 

PAPAVEEACEiE, 99, 500 

Papilionacea, 397, 502, see Legumi- 

Paris quadri folia, 31, 358, 360, 367, 396 
Paritium, 137 
Parnassia, 364 
Paronychia serpyllifolia, 473 
Parthenium iuodoruro, 243 
Passiflora, 137, 201, 301, 365, 463, 473 

ccerulea, 185 

pahnata, 185 

quadrangularis, 181 


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 

grandiflorura, 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 
Phalamopsis, 238 

amabilis, 227 

equestris, 231, 238 

Schilleriana, 224 
Pharbitis, 73 
Phaseolus, 21, 298, 364 
Philadelphacea, 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, 7:? 

Phoenii dactylifera, 318 

Phylica, 326 

Phyllanthua longifoliuB, 198 

Phyllartbron, 64, 396 

Phyteuma odoratum, 271 

orbiculare, 20 

Bpical inn, 16<>. 252 
Phytolacca, 20 
Picea, sou Abies 

Lowii, 203 

nobilis, 203 

Nordmanniana, 203 

Webbiana, 203 
Picrasma, 509 
Pimpinella magna, 66 

Saxii'ragra, 66 
Pinckneya, 120 

Pinna, 11, 19, 114, 31S, 346, 471, (see 
Abies, Larix, Cedrus) 

alba, 192 

pinea, 90, 218 

Pinaster, 13, 21 

sylvestris, 21, 349 
Piatacia Lentiscus, 197 
Pisum, 137. 295 

sativum, 27, 30, 31, 95, 166, 302, 
372, 432, 172, 502, 510 
Platycodon, 296 

grandiflorum, 504 


Plantago, 114, 136 

Coronopus, 458 

lanceolata, 61, 104, 108, 110, 111, 

media, 20, 111 

major, 65, 108, 109, 111, 243, 372, 

maritima, 108 
Platanus, 50 

Plectranthus fruticosns, 238 
Pltjmbaginace.e, 114 
Plumbago, 360 
Poa alpina, 169 

annua, 169 

bulbosa, 169 

pratensis, 169 

trivialis, 169 . 
Podalyria myrtillifolia, 44 
Podocarpna, 426 
Podophyllum, 295 

peltatum, 500 
Podospermum laeiniatum, 247, 250 
Poggendorffia, 294 
Pogouia ophioglossoides, 23S, 386 
Poinsettia, 340 
Polemonium caeruleum, 20, 66, 253, 

305, 310, 404 
Polianthea, 296 

Polygala vnlgarU, 399 

POLYGONAOB B, I I I. L38, 150 

Polj gonatnm ancepa, 12, L5 

mnltifolium, -■'>, -W. 
Polygonum, 1 1 I, 365 
' orientate, 31, 390 

viviparum, 106, 169 
Polypodium anomalnm, 190 
Pomack.v:, 70, 71, 77, 79, 142, 405, 

Pomaderria elliptica, 355 
Pomax, 38 
Populua, 202, 309 

alba. 66 
Portnlaca, 295 
Potamogeton, 329 

bifolinm, 434 
Potentilla, 139,140, 149, 295, 371. 138 

alpestris, 503 

argentea, 280 

auseriua, 503 

major, 374 

nepalensis, 250, 252, 275, 280 

reptans, 503 

Tormentilla, 503 
Poterium, 113 

Sangniaorba, 107 

polygamum, 281 
Primula, 71, 73, 114, 121, 138, 150, 
296, 335, 360, 365, 367 

acaulis (see vulgaris), 15, 2 18, 250, 
308, 310, 377, 504 

Auricula, 45, 133, 271, 282, 504 

denticnlata, 20, 504 

elatior, 45, 250, 504 

calycantbema, 283 

imperialis, 103 

officinalis, 250, 283 

prsenitens, 271, 504 (see sinensis) 

sinensis, 24, 45, 95, 98, 103, 133, 
253, 2.36, 262, 263,271, 281, 297, 
314, 315, 3S9, 429, 441, 449 

variabilis, 104 

veris, 17, 20, 35, 39, 45, 105, 164 

villosa, 504 

vulgaris, 17, 70, 106 (see acaulis 
and officinalis) 
Pkimulaceje, 98, 114, 118, 121, 133, 

138, 150, 262, 339, 366, 504 
Prismatocarpua, 114, 150, 178 
Prunella vulgaris, 226 
Prunua, 137, 203, 295, 300, 360, 361, 
367 (see Cerasus, Amygdalus, 


Armeniaca, 44 

Cerasus, 7 1 

domestica, 366, 464 

Lauro Cerasus, 21, 446 

apinosa, 44 



i.\i»r;x or names of plants. 

Prunus sylvestris, 21 
Pseudostachyum polymorphmn, 168 
Psittacanthus, 371 
Ftelea, 36-4 
Pterandra, 290 
Pteridopliyllum, 331 
Pterisanthes, 328 
Pteris quadriaurita, 333 
Puuica Granatum, 21, 295, 325 
Fyvethrum, 114 

iuodoruin, 131 

Parthenium, 339 
Pyrus, 82, 107, 113, 121, 137, 149, 203 

communis (Pear), 36, 66, 162, 178, 
422, 423, 503 

dioica, 304 

japonica, 161 

Mains (Apple), 44, 78, 79, 166, 2-M>, 
250, 282, 295, 310, 325, 327, 375, 
-106, 420, 503 

spectabilis, 508 

torminalis, 325 


' Juercus, 51 
Cerris, 66 
pubescens. 66 


Raphanus sativus, 161, 252, 327,360, 

RAMTTNCtTLACEJE, 113, 122, 136, 148, 

195, 246, 339, 410,499 
Ranunculus, 19, 55, 113, 119, 258, 288, 
2H5, 296, 297, 360, 367, 407 

aconitifolius, 500 

acris, 67, 250, 436, 500 

aqnatilis, 458 

asiaticns, 500 

auriconras, 195, 285, 397, 406, 461, 

bulbosas, 17, 20, II, 195, 106, 500 

bullatus, 500 

Ficaria, 70, 96, 195, 261, 368, 106 

flnitans, 433, 437 

gramineus, 500 

Lingua, 44, 67 

Philonotis, 20, 339 

tripartitus, 20, 46 

repens, 23, 252, 261, 500 
Kesedace^, 137, 148 
Reseda, 67, 137, 141, 148 

lutea, 252, 271 

Lnteola, 61, 112, 399 

odorata, 20, 11, 2 

Pbytenma, 252, 261 
Restiace i . 1 L5, 167, 189, 350, 463 

Restio, 115, 194 
Rbamnns catharticus, 463 

Frangula, 280 
Rliinuntbus crista galli, 238 
Rhodiola rosea, 197 
Rhodora, 73 

Rhododendron, 72, 73, 91, 133, 137, 
176, 289, 290, 296, 453 

indicum, 504 

linearllobum, 72 
Rhus Cotinus, 62, 66, 348, 409, 467, 

Toxicodendron, 66 
Ribes, 296 

nigrum, 66, 296 

sanguineum, 503 
Richardia tethiopica, 337, 358 
Ricinus communis, 198, 300 
Ricotiaua, 364 
Robinia, 365 

Pseudacacia, 44, 204, 317, 325, 396 
Rosacea, 70, 71,77, SO, 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 

nis'ea, 502 

pimpmellifolia, 5<i2 

rubiginosa, 502 

spinosissima, 502 

sulphurea, 502 
RUBIACE-E, 45, 80 
Rubia tinetorum, 322, 32S 
Rulms, 137, 252, 279, 399. 106, 429, 

arcticus, 67 

cajsius, 281, 285, 502 

corylifolius, 502 

fruticosus, 66, 281, 374, 380, 461 

Idjeus, 396 

rosifolins, 502 
Rudbeckia, 82, 111, 127 



Uiiellia rlaticlc-tina, 108, K) I 

Komex, r>. 138, L60, 825 
uifolioa, -'7--. 281 
criapns, 804, 810, 1 is 
- bates, 878, 281, 131 

Rosens, 828, 170 

aculeatns, 60, 318, 3ls, 115 
Ilia jnncea, -" 

RUTACU, 187, 1 19 

lluta, 363, 367, 371 

Sagina, 307 

procumbens, 501 
Sagittaria, 296, 329 

latifoliu. ■" 7 

dtolia, 325. 5^7 
Salisbnria adiantifolia, 61 
Salix, 166, 16s, 244, 310, 326, 349 

babvlonica, 202,257,262, : 

calycnlata, 29 

eaprsea, 271, 299 

cinerea, 41, 45, 189, 299 

fragilis, 357 

monandra, 29 

nigricans, 299 

pendula, 357 

repens. 191 

silesiaca, _ 

vitellina, 21 
Salpislossis straminea, 41 
Salvia, 226 

officinalis, 66 

pratensis, 36 

Yerbenaca. >7 
Sambucus, 296, 359, 360, 36S 

nigra, 21, 55, 66, 164, 325, 396 

raceuaosa, 66 
Sanguinaria, 295 

canadensis, 500 
Sanguisorba, 113 

officinalis, 107 
savtalaceje, 122, 138, 150 
Sapi>-dace.e, 137, 448 
Saponaria, 28, 67, 76, 149, 295. 297, 

300, 360, 379 

officinalis, 97, 301, 501 
Sarothamnus, 360, 363 

Scoparins, 363 
Saxifraga, 33, 44, 106, 113, 296, 390 

crassifolia, 307, 310 

decipiens, 288 

foliosa, 2S1 

granulata, 298, 504 


longitolin, 404 

Saxifraga luutata, _'' i 
Saxifuagac e i . 80, si 

15, L06, 11 1, 138, 1 11, 296, 

agrestia, 252 

arvensia, 325 

atropnrpnrea, 62, 66 

Columbaria, 27,2, 2->l 
Scandix, 125 

Schismatopera distichophylla, 191 
Schcenodorus, 364 adnot 
Schcenus oephalot< - 
Schoenodon, 365 
Sciadopitvs verticillata, 352 
Scilla, 296 

autumnal'-, 5l ■' 

nutans, 507 
Scirpns lacnstris, 325 
Scolopendrium D'Urvillei, 332 

vulgare, 64, 311,326, 328, 159, i67 
Scorzonera, 19 

octangularis, 247, 250 
Scropbularia, 114, 455, 463 

aqnatica, 226, 238, 262, 281 

arguta, 334 

nodosa, 44, 106, 281 
Sceophulaeiacej:, 114, 121, 137, 150, 

222, 505 
Secale cereale, 55, 115 
Sechinm edule, 81, 21 «2 
Sedum cristatnm, 18, 20 

reflesnm, 20 
Selenipedium caudatum, 224, 227 
Selinnm, 82, 114 

caruifolmm. 281 
Sempervivuiri, 105, 113, 204, 509 

montanuin, 310 

sediforme, 164 

tectornm, 292 adnot, 308, 310 
Senecio, 44, 114 

vulgaris, 247, 250, 252, 339, 432, 
437. 439. 457 

-.., 296, 377, 504 
Sesamum, 365 

indicum. 23S 
Seseli, 113, 281 

coloraturu, 166 
Sideritis, 238, 365 
Silaus, 113, 365 
Silene, 44, 137. 148, 295, 301 

conica. 68 

Otites, 464 

indata, 501 
Sinapis, 57, 136, 270, 295, 371 

arvensis, 181, 250, 261, 264. 271, 
280, 501 
Sinningia purpurea, 431 
Sisymbrium, 136 

officinale, 250, 261, 271, -- 



Sisymbrium tenuifolium, 252 
Slateria, 74 

SOLANACE35, 150, 430, 505 
Solarium, 73, 125, 150, 296, 360, 365, 
367, 370, 424, 453 

amazonicum, 430 

Dulcamara, 66, 288, 430, 504 

esculenturn, 74 

cornutum, 430 

Lycopersicuin, 38, 44, 74, 171, 391, 

tridy natrium, 430 

tuberosum, 54, 142, 288, 333, 420 

Vespertilio, 430 
Sophora, 55 
Spartiantbus, 295 

junceus, 502 
Spartium, 295 

junceum, 18, 21 

Scoparium, 21 
Spathiphyllum, 245 
Spilanthes, 138, 365 

oleracea, 44 
Spinacia oleracea, 31, 197 
Spira?a, 21, 137, 295 

Filipeudula, 503 

oblongifolia, 252 

primifolia, 503 

Reevesii, 503 

strobilacea, 503 

Ulmaria, 503 
Spiranthes, 319 
Splacbnum vasculosum, 49 
Stachys, 138, 339, adnot 

germauica, 310 

lanata, 375 

sylvatica, 74, 253, 271, 281 
Stackhousia juncea, 334 
Stangeria paradoxa, 172 
Stapelia, 20 
Stapbylea pinnata, 30 
Stauntonia latifolia, 194 
Stellaria, 148, 404, 406 

media, 271, 279, 280 
Stellate, 396, 408 
Steeculiace^;, 362, 464 
Sterculia platanifolia, 20, 256 
Sternbergia, 296 

lutea, 506 
Stratiotes aloides, 406 
Strelitzia juncea, 459 

regina, 25 
Streptocarpus Rexii, 15, 44, 226, 227, 

Strophantus, 326 
Suaeda, 365 

fruticosa, 430 

maritima, 20, 21 
Suregada, 198 

Sympboricarpus racemosus, G6 
Symphyomyrtus, 38 
Symphytum, 71, 138, 365 

officinale, 253, 262, 263, 271 

Zeyheri, 271 
Syringa, 296, 360, 367 

persica, 44, 61, 66, 284, 505 

vulgaris, 76, 505 


Taberna3montana, 296 

coronaria, 504 
Tacsonia pinnatistipula, 294 
Tamus communis, 21 
Taraxacum, 164 (see Leontodoii) 
Taxodium, 114 

disticbum, 444 
Taxus baccata, 11, 21, 90 
Tetragonia, 113 

expansa, 142, 180 
Tethagoniace^;, 149 
Teucrium campanulatum, 226, 233, 238 
Tbalictrum, 295, 500 

minus, 307, 310 
Thea, 295, 502 
Thelymitra, 224 
Thesium, 121, 123, 138, 150 
Tblaspi 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 

europtea, 22, 30, 65 

parvifolia, 66 
Tiliacea?, 137, 362 
Tithonia, 26 
Tofieldia, 296 

calyculata, 300 
Torenia scabra, 406 
Torilis, 82, 149 

Anthriscus, 256, 271, 281, 339, 
406, 441, 4 12 
Tradescantia, 245, 296, 360, 365, 367, 

virginica, 88, 507 
Tragopogon, 19, 20, 73, 114, 431 

orientale, 250 

pratense, 217, 2Z.0, 442, 457 
Trapa nutans, 202, '■'•('• 1 
Tricliostomuni rigidulum, 49 
Trifolium, 106, 113,121, 137,139, 1 49. 
295, 360, 364 



Trifolinm hybridum, 262, 899, 106, 
pretense, 2o 

ivpens, 20, 23, OS, 70, 98, 145, 1 16, 
226, 231, 238, 250, 252 256, 
260, 262, - ,;: >> 271, 276, 27'.', 
-. 397, 399, 406, 107, 
l:i!. 136, 138,502 
resupinatnm, 20 
Trillium grendinorum, 507. 510 
Trinia vulgaris, IM."> 
Triphasia aurantiaca, 369 
I'riti.uni, 86, 115, 350, 391 
repens, 106, 325 
vnlgare, 55, 113 
Triumt'etta, 137, 252, 259, 260, 262, 

265, 271, 280 
Trollius europaeus, 66, 295, 500 
Tropj.oi.acej:, 119, 501 
Tropseolnm, 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, 

svlvestris, 507 
Turritis, 271 

glabra, 252 
Typl.a, 189 


Hex, 295, 360 

europaeus, 377, 502 

Ulmus americana, 66 

campestris, 31, 52, 62, 157, 325, 
353, 427 

Umbellifeej., 37, 71,73,80,107,113, 
121, 127, 131, 132, 137, 139, 140, 
143, 149, 150, 162, 244, 257, 261, 
339, 368, 395, 405, 406, 407. 437, 

Oiedo Candida, 279 

I'bticace j., 114 

Urtica dioica, 62, m, 191 

Usteria, 283 

Valeriana, 11 1, If',:,, 119 

dioica, 325 

officinalis, 325 
Valisneria spiralis, 319, 133 
Verbascnm, 7:!, 116, L37, 225, 253, 13 I 

anstrale, 35 

Digram, 22i>, 251 

pblomoides, 281 

Tbapsns, In 1 
Verbena, 67, 68, 506 
Veronica, 60, 150,296, 331, 360, 361, 
365, 375, 505 

austriaca, 66 

Beccabunga, 468 

Chamsedrys, 1 12 

latifolia, 459 

longifolia, 325 

spicata, 325, 374 
Viburnum, 44, 296, 417 

Opulus, 473 
Vicia, 30, 365. 369, 172 

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, 

grandiflora, 501 

hirta, 226, 238 

odorata, 20, 94, 220, 226, 238, 286, 
404, 429, 501, 508 

tricolor, 461, 501 
Violace.e, 137, 334, 403, 406, 500 
Viscum album, 56, 369, 509 

VlTACE J., 137 

Vitex incisa, 238 
Vitis, 417 

vinifera, 20, 29, 34, 44, 66, 137, 157, 
182, 183, 211,280, 374, 422 424 


Wedelia perfoliata, 442 
Weigela rosea, 358 
Welwitechia, 162, adnot 
Wildenovia, 115, 167, 168 
Wistaria, 364 
sinensis, 226 


Vaccinium, 68 
Valantia cruciata, 44 
Valeriaxace.e, 114 

Xanthosoma appendiculatum, 31 
Xanthoxylum, 21 
Xvlophylla, 328 





Yucca, 361 

tiexilis, 361 

Zamia, 170 

Zea Mays, 21, 113, 136, 191, 197, 300, 

310, 350, 369, 466 
Zingiber Zerumbet, 224, 227 
Zinnia, 4 1 

elegans, 20 
Zygopetalum maxillare, 398 


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 Sempervivvn tecot<rmvi read Semper- 
vivum tectorum. 




Made In Italy 

N*m Yort Botanical O*ar0*n Lito*a»» 

QK664 M37 9*n 

Masters. Maxwell Ty/Vegetable teratology _ 

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