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VENDU EN 1922

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INSTITUTED MDCCCXLIV.

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This volume is issued to the Subscribers to the Ray

the Year 1868.

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VEGETABLE TERATOLOGY,

AN ACCOUNT

OF THE

PRINCIPAL DEVIATIONS FROM THE USUAL
CONSTRUCTION OF PLANTS

BY

MAXWELL T. MASTERS, M.D., F.L.S.

LIBRARY
NEW YORK
BOTANICAL

GARDEN

WITH NUMEROUS ILLUSTRATIONS BY

E. M. WILLIAMS.

. : L. OEE ON:
PUBLISHED FOR THE RAY SOCIETY BY
ROBERT HARDWICKE, 192, PICCADILLY.

MDCCCIXIX.

TO

JOSEPH DALTON HOOKER, M.D.,

D.C.L., LL.D., F.R.S., F.L.S., ETC. ETC.
DIRECTOR OF THE ROYAL GARDENS, KEW,
This Volume

Is

GRATEFULLY INSCRIBED.

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AUG 7 - 1923

LIBRARY

NEW YOR!
BOTANICAL
GARDEN
TABLE OF CONTENTS.
INT EOVUCTEO N xxi
ib OOK Ul:

DEVIATIONS FROM ORDINARY ARRANGEMENT

PART I.—Union or OrGaAns

CEVA DBE:
COHESION

Cohesion between axes of same plant, 9—Fasciation, 11—
Cohesion of foliar organs, 21—Of margins of single organs, 21
—Tubular petals, 23—Cohesion of several organs by their
margins, 25—Of the sepals, 27—-Of the petals, 28—Of the
stamens, 29—Of the pistils, 29— Ascidia or pitchers, 30.

CHAPTER IL.
ADHESION

Adhesion of foliar organs, 32—Of leaves by their “surfaces,
33—Of foliar to axile organs, 34—Of sepals to petals, 34—Of
stamens to petals, 34—Of stamens to pistils, 35—Miscellaneous
adhesions, 35—Of fruit to branch, 36—Synanthy, 36—Syn-
carpy. 45—Synspermy, 50—Between axes of different plants
of same species, 50— And of different species, 55—Synophty, 56.

32

vill TABLE OF CONTENTS.

PAGE
PART II.—InDEPENDENCE or ORGANS - 58
CHAPTER I.

FISSION . ; 2 : F : . 59
Fission of axile organs, 60—Of foliar organs, 61—Of petals,

66—Of stamens, 68—Of carpels, 68.
CHAPTER II.
DIALYSIS 69

Dialysis of margins of individual parts, 70—Of margins of
parts of same whorl-calyx, 70—Of corolla, 71—Of stamens,
73—Of carpels, 73.

CHAPTER Iii.
SOLUTION

Solution of calyx from ovary, 77—Of stamens from petals,
82.

PART II].—Atrerations or Position

CHAPTER I.
DISPLACEMENT ; : . s :
Displacement of bulbs, 84—Of inflorescence, 85—Of leaves,

87--Of parts of flowers, 91—Of carpels, 96—Of placentas and
ovules, 96.

CHAPTER II.
PROLIFICATION

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—Avxillary prolification, 138—Foliar, 141—Floral, 142
—Complicated prolification, 151—Of embryo, 155.

76

83

84

100

TABLE OF CONTENTS. 1X

CHAPTER III.

HETEROTAXY , F : : ; *156

Formation of adventitious roots, 156—Of shoots below the
cotyledons, 161—Adventitious leaves, 162—On scapes, 163—
Production of leaves or scales in place of flower-buds, 164—
Viviparous plants, 168—Formation of buds on leaves, 170—In
pith, 171—On bulbs, 172—Production of gemme 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 Composite, 188—Heterotaxy affecting the inflorescence, 188
—Supra-soriferous ferns, 190.

CHAPTER IV.
HETEROGAMY ‘ F 5 : : . 190
Change in the position of male and female flowers, 191—From
monecious to dicecious condition, 193—From diccious to
moneecious, 193—From hermaphroditism to unisexuality, 195—
From unisexuality to hermaphroditism, 197—Pollen replaced
by ovules, 201.

CHAPTER V.
ALTERATIONS IN THE DIRECTION OF ORGANS 5 . 201

Fastigiation, 202—Eversion, 204—Altered direction of leaves,
205 — Altered direction of flower, 206—Reflexion, 209 --
Gymnaxony, 211.

BOOK. EE
DEVIATIONS FROM ORDINARY FORM ; PAB:
PART 1I.—Srtastmorpuy 4 : : . “216
CHAPTER I.
PERSISTENCE OF JUVENILE FORMS. SPAY

Stasimorphy in leaves of Conifers, 217-—Regulax ee 219.

x TABLE OF CONTENTS.

PART I1.—P.Letomoreny

CHAPTER I.
IRREGULAR PELORIA

PART III.—Meramorreuy

CHAPTER I.
PHYLLODY

PAGE

. 228

228

241

Phyllody of bracts, 242—In inflorescence of Conifers, 245—
Of calyx, 245—Of corolla, 251—Of stamens, 253—Of pistils, 256
—Of ovules, 262—Changes in nucleus of ovule, 269—Phyllody
of accessory organs, 272—-Chloranthy, 273—General remarks
on, 278.

CHAPTER IT.

METAMORPHY OF FLORAL ORGANS .

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.—HetTERomorrPHyY

CHAPTER I.

DEFORMITIES

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.

TABLE OF CONTENTS. x]

CHAPTER II.

PAGE
POLYMORPHY : ; : . 3829
Heterophylly, 330—Dimorphism, 333—Sports or bud-varia-
tions, 336.
CHAPTER III.
ALTERATIONS OF COLOUR . ; : : . 3837

Albinism, 337— Virescence, 3388—Chromatism, 339.

POO KE rE
DEVIATIONS FROM ORDINARY NUMBER ‘ . 340
PART I.—Increasep NuMBER oF ORGANS f . 343
CHAPTER I.
MULTIPLICATION OF AXILE ORGANS—INFLORESCENCE . 846

Multiplication of branches, 346—Plica, 346—Polyclady, 347
—Multiplication of branches of inflorescence, 348—Of bulbs,
350—Of florets, 351.

CHAPTER II.
MULTIPLICATION OF FOLIAR ORGANS ; . eS uy

Pleiophylly, 8353—Multiplication of stipules, bracts, &e., 357
—Polyphylly, 358—Increased number of leaves in a whorl, 358
—Polyphylly of bracts, 358—Of calyx, 358—Of corolla, 359—
Of andrecium, 361—Of gynecium, 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—Andrecium, 379—Andreecium
of Orchids, 380—Pleiotaxy of gynecium, 388—Increased num-
ber of flowers in an inflorescence, 391.

xu TABLE OF CONTENT'S.

PAGE

PART II.—Diminisoep NumBer or Orcans _ 392
CHAPTER I.
SUPPRESSION OF AXILE ORGANS P : . 393

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

CHAPTER II.
SUPPRESSION OF FOLIAR ORGANS . : e . 395
Aphylly, 395—Meiophylly, 396—Of calyx or perianth, 396—
Of corolla, 397—Of andrecium, 398—Of gyneecium, 399—Of
flower, 400—Meiotaxy, 403—Of calyx, 403—Of corolla, 403—
Of andrecium, 405—Of gynecium, 406—Of ovules or seeds, 407
Of flower, 408— General remarks on suppression, 409.

BOOK IV.
DEVIATIONS FROM ORDINARY SIZE AND CONSIST-
ENCE : : : ; ; . 413
PART [.—Hyrerrtrorny é . 416
CHAPTER I.
ENLARGEMENT. 4 P Bikey

Of axile organs, 418—Knaurs, 419—Hnlargement 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 andreecium, 430—Of gyncecium,
430—Of fruit, 431 —Alterations of consistence, 432.

TABLE OF OONTENTS. Xill

CHAPTER II.
PAGE
ELONGATION ; ; 5 : . 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.
CHAPTER III.
ENATION : 7 : : . 443

Excrescences from axile organs, warts, 444—Enation from
leaves, 445—From sepals, 448—From petals, 448—Catacorolla,
451—Enation from stamens, 453—From carpels, 453.

PART II.—Arrorny ; : . 454

CHAPTER I.
ABORTION : ; : : ; . 455

Abortion of axile organs, 455—Of receptacle, 457—Of leaves,
458—Of perianth, calyx, and corolla, 460—Of stamens, 463—
Of pollen, 463—Of pistil, fruit, &c., 464—Of ovules, 466—De-
pauperated ferns, 466—General remarks, 467.

CHAPTER II.
DEGENERATION . : : . 470

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

GENERAL CONCLUSIONS.

General morphology of the leaf and axis, 476. Homology.
476—Special morphology, 479—Calyx-tube, 480— Andreecium,
482—Inferior ovary, 482—Placentation, 483—Structure of the

X1V TABLE OF CONTENTS.

: PAGE
ovule, 484—Leaves of Conifers, 484—Relative position of

organs, 484—Law of alternation, 485—Corelation, 486—Com-
pensation, 488—Teratology and classification, 488.

APPENDIX.

Double flowers, varieties of, 491—Causes of production, 491—
Relation to variegated foliage, 497—List of plants producing,
499.

NOTE. : : ; : J . 508
INDEX TO SUBJECTS . : : ,; . dll

INDEX OF NAMES OF PLANTS . ; : . 517

FIG.

go fo

oo

eS) eI

10.

a
12.
13.
14.
15,
U7.
18.

19,
21.

LIST OF ILLUSTRATIONS.

PAGE

Diagram of regular pentamerous flower 4
Cohesion of branch Dipsacus sylvestris 10
Fasciated lettuce (‘ Gard. Chron.’) 11
Fasciation in Asparagus 12
5 in Pinus Pinaster 15

and spiral torsion in Asparagus ( Gard. Grea 9) 14

5 in scape of dandelion 16
Pitcher on leaf of Pelargoniwm 22

Transition from flat to tubular segments of the perianth in
Eranthis ¢ :
Pitcher of Crassula ar Beeaans (C. Morren)
Gamopetalous corolla, Papaver bracteatum . :
Adhesion of petaloid stamen to segment of perianth, Crocus
of petal, stamen and carpel, Cheiranthus Cheiri
Synanthy, Campanula Mediwn (‘ Gard. Chron.’)
16. Synanthic flowers of Calanthe vestita
Synanthy in Digitalis purpurea (‘ Gard. Chron.’)
= in Calceolaria
20. Syncarpic apples
Adhesion of two stems of oak (‘ indi Bheon )
of branches of elm (‘ Gard. Chron.’)
a of two roots of carrot, one white, the other ba
(‘ Gard. Chron.”’) , .
Section through inverted and adherent mushrooms .

). Bifureated male catkin, Cedrus Libani

PA leaf, Lamium album

LIST OF ILLUSTRATIONS.

PAGE
. Bifurcated leaf, Pelargonium ; : ; . 63
Ps frond, Scolopendrium . : 2 . 64
. Three-lobed lip of Oncidiwm . 68
. Dialysis of corolla in Correa Th
5 of corolla in Campanula (De Candolle) 72
2. Anomalous form of orange 74
34. Disjoined carpels of orange (Maout) 75
. Proliferous rose (Bell Salter) 78
37. Apple flower, with detached calyx, &e. : 79
. Flower of Gnanthe crocata, with detached calyx, &c. . 80
. Anomalous bulbs of tulip : J . 85
. Displaced leaf of Gesnera (C. Morren) : : . SS
. Leaves of Pinus pinea . : ; : . 89
. Deranged leaves of yew . 90
. Cohesion of sepals and displameniGn of nit of ‘Oneida
cucullatum . : : . 92
. Malformed flower of Cuptnedinnd' 27: : , ee
. Diagram ofthe same . : 5 . 93
. Of natural structure in oo : . 8S
. of malformed flower of Lycaste Skinneri . 3)
na of malformed flower of Dendrobium nobile . . 94
. Natural arrangement in same flower ; . 94
. Diagram of malformed violet : ‘ . 94
. Monstrous flower of Cerastiwm (‘ Gard. Chron, ) : 97
. Inflorescence of Polyanthus, with tufts of leaves at the summit ; 105
. Lateral prolification in inflorescence of Pelargoniwm . 108
. Paniculate inflorescence of Plantago major . : . 109
. Branched spike and leafy bracts of the same ; . 110
. Inflorescence of Plantago lanceolata, with leaves and secondary
flower-stalks at its summit. : . ill
. Branched inflorescence of Reseda Iuteola . : . 112
. Thalamus of strawberry prolonged into a leafy branch (‘Ameri-
can Agriculturist ’) ; . 116
. Flower of Verbascwm, with dialysis of ais and pee and
prolonged thalamus . : 2 . 116
. Median floral prolification of Dianthus : ; . 120
. Leafy carpels and prolification of Daucus Carota . 123

. Median floral prolification of Delphinium . ; . 126

LIST OF ILLUSTRATIONS, XVll

FIG. PAGE
63. Median prolification, &c., of Orehis pyramidalis i . 128
64, Proliferous rose (Bell Salter) _ . 130
65. Axillary floral prolification of Nymphea Sain (‘ ied: Chron.”) 144
66, 5 3 of Dianthus ' , . 146
67. Proliferous rose - d : . 151
68. Proliferous rose (‘ Gard. es se : . . 152
69. Diagram of prolified orchid : : : . 153
70. 7 - orchis ; : . . 154
71. Adventitious roots from petiole of celery . : . 158
72. Germinating plant of mango ; : . 159
73, 74. Adventitious roots from leaves : : . 160
75. Hip of rose bearing leaf 2 . 162
76. Leaves proceeding from the ovary of igi | . 162
77. Flower-stalk of dandelion, with leaves ? . . 163
78. Tuft of leaves in place of flowers in Valeriana sp... . 165
79. Scale-bearing spikelets of Willdenovia ; : . 167
80. “Rose Willow ” ; ; : . 167
81. Viviparous flowers of Aira vivipara , : . 169
82. Formation of shoot on leaf of Episcia bicolor : ep
83. Adventitious buds on root of sea-kale ; Bee
84,85. Production of adventitious bulbs in hyacini h (* Gard.

Chron.’) . ; ; . 172
86. Adventitious buds on hyacinth (‘ Gard. oe a} : . 173
87—92. Nepaul barley 2 : ; 174, 175
93. Formation of buds on fruit of fests : : - ES
94. Flower-bud in the pod of Sinapis d een
94*, 95. Adventitious pod in silique of Chetrintheas : . 182
96, 97. Grapes, with adventitious fruits in interior : Bes fo
98. Stamens in ovary of Beckea diosmifolia . : . 184
99. Pollen in ovule of passion flower (S. J. Salter) : . 185

100. Female flowers at the summit of the inflorescence of Carex

acuta f : F >, ee

101. Monecious hop (‘ Gard. ‘Gison: ). : : . 193

102. Superior ovary, &c., of fuchsia. : : . 198

103. Hermaphrodite flower of Carica . : . .. 199

104. Ovuliferous anthers of Cucurbita sp. ; ie . 200

105, 106. Prolonged inflorescence of fig. ; : . 205

107. Hollow turnip, with inverted leaves : 4 . 206

xVill LIST OF ILLUSTRATIONS.

FIG, PAGE
108. Normal flower of Glowinia : ; : . 207
109. Regular peloria of Glowinia ; : . 207
110, 111. Structural details of erect Gloxinia . 5 . 208
112. Reflected corolla of azalea F . 209
113, 114. Enlarged and erect placenta of Gaia anime (C.
Morren) . : . : , . 210
115. Dimorphic leaves of Juniperus sinensis : ; 227
116. Regular peloria of Delphinium . : : . 219
117. Structural details of the preceding ; , . 219
118. Regular peloria of violet : ; . 220
119. Double-flowered regular violet 220
120. Regular peloria of Eccremocarpus scaber_ . ‘ . 222
1174] rs Pe Cattleya : F - 223
122. Peloria of Calceolaria . ; : . 230
123. Peloric flower of Aristolochia : ; .. 232
124, 125. Peloric flowers of Corydalis . : . 236
126. Rose plantain . ‘ : . 242
127. Leafy bracts in Beaders maj ‘ : .. 248
128. ,, scales of Dahlia . 4 : . . 244,
129. ,, sepals of rose : : F P . 246
UBUy is Fuchsia (‘Gard. Chron.’) . : . 247
131. ,, calyx of primrose ; ; : . 248
W32.- 5 As melon . . ¥ : . 248
133. ,, sepals and petals of Geraniwm ; : . 251
134. ,, stamens of Petunia : ‘ . 254
135, 136. Leafy anthers of Jatropha (Miller) d : . 255
137. Proliferous rose, with leafy carpels, &c. (Bell Salter) . 207
138. Cucumber, with adventitious leaf attached (8. J. Salter) . 258
139. Leafy carpels in flower of Triwmfetta 5 ; . 260
140. ,, ovules of Sinapis. j d . 264
141, 142. Leafy ovules of Trifoliwm repens (Gaza y) : . 265

143. Portion of leafy carpel of Delphinium, with ovules (Cramer) . 266
144, 145. Enlarged view of section of leafy carpel, &c., of Del-

phinium (Cramer) : : . 267

- 146. Placentz of Dianthus, bearing ovules se waned (‘ Gard.
Chron.’) . . 268
147. Ovules passing into daapers Dianthus (‘ Gama: Gea a) . 268

148. Leafy shoot in place of ovule of Gaillardia . . 270

Lia.

LIST OF ILLUSTRATIONS.

149, 150. Leafy sepals, petals, &ec., of Epilobium .

151.
152.
153.
154.
155.
156.
157.
158.
159.
160.
161.
162.
163.
164,
165.
166.
167.
168.
169.
170.
171.
172.
173.
174.

175.

176.
177.
178.

Leafy carpel of rose, with deformed ovules .
Flower of St. Valery apple
Petaloid calyx of Mimulus
Double stellate columbine
Four-winged filaments of Rhododendron
Diagram of malformed flower of Catasetwm .
Petaloid stamen of Viola
Double columbine
Petaloid stamens of Hibiscus
Displaced coloured leaf, &c., of tulip
Supernumerary carpels in orange (Maout) .
Pistilloid stamens of poppy

Y is wallflower, &ec.
Passage of stamen to carpel in lily
Transition of stamens to carpels, Sempervivum tectorwm
Ascidia of cabbage
Stalked pitcher on lettuce leaf
Tubular petal of Primula sinensis .
Spurs on flower of Calceolaria
Contorted stem of Juncus

3 branch of Crategus
Spirally-twisted stem of teazel

5 E Galium .
i root (‘ Gard. Chron.’)
Interrupted growth of radish (American Agricultur ist)
” » inapple

Polymorphous leaves of lilac ;
Adventitious growth on frond of Pteris quadriaurita .

. 30d

179, 180. Coloured flower-stalks of feather hyacinth (C. Morren) 347, 348

181.
182.

183,
185.
186. Multiplication of parts of flower in a plum ¢ Gard. Cheon )

187.
188.

189.

Multiplication of catkins, Corylus .
Branched inflorescence of brocoli (‘ Gard. Chiat 2)
184, Supernumerary leaf of elm .

” - hazel

Wheat-ear carnation
Multiplication of bracts in Delphiinivom Gonasiinn
” rp Pelargonium

. 349
. 351

303, 354

. 855
. 366
. 372
. 373
. 373

LIST OF JLLUSTRATIONS,.

. Double white lily
. Double flower of Campanula seiussagietin :
. Diagram of usual arrangement of parts in Orchis (Darwin)

a malformed flower of Ophrys aranifera

. Malformed flower of Ophrys aranifera ;

. Diagram of malformed flower of Orchis mascula (Crawier)
. Multiplication of carpels, Tulip

. Section of St. Valery apple

. Regular dimerous flower of Calanthe eealits

3 = Odontoglossum Alexandre

. Hypertrophied branch of Pelargonium
. Tubers in the axils of leaves of the potato .
. Hypertrophied pedicels of ash

204. Hypertrophy and elongation of dona: Sate &e., in

pears

. Hypertrophied pinian the Cocos re

. Elongation of flower-stalk, Ranunculus acris

. Linear leaf-lobes of parsley ;

. Passage of pinnate to palmate leaves in es chesnut
. Elongation of thalamus, apostasis, &c., in flower of Delphinium

(Cramer)

. Adventitious growths from aden leaf
. Crested fronds of Nephrodium molle
. Supernumerary petals, &c., Datura fastuosa

5 petaloid segments in flower of Gloxinia

. Catacorolla of Gloxinia (E. Morren)
. Atrophied leaves of cabbage

. Abortion of petals, pansy

. Flower of Oncidium abortivum

. Bladder plum .

422, 423

INTRODUCTION.

Tritt 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
sahent 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. Linnzeus 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

XXll INTRODUCTION.

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
observers.

While the number of scattered papers on vegetable
teratology in various European languages is so great
as to preclude the possibility of collatimg them all,
there is no general treatise on the subject in the

INTRODUCTION. Xxill

English language, with the exception of Hopkirk’s
‘Flora Anomala,’ a book now rarely met with, and
withal very imperfect; and this notwithstanding that
Robert Brown early lent his sanction to the doctrines
of Goethe, and himself illustrated them by teratological
observations. In France, besides important papers of
Turpin, Geoffroy de Saint Hilaire, Brongniart, 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, Jeeger, 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 ‘ Eléments,’ to be
considered as referring to limited portions only and
not to the whole subject.’

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 in the various
European scientific publications have also been laid
under contribution. In most cases reference has been
given to, and due acknowledgment made of, the sources
whence information has been gathered. Should any
such reference be omitted, the neglect must be attri-
buted to imadvertence, not to design. Im selecting

' An excellent summary of the history of Vegetable Teratology is

given in Kirschleger’s ‘ Essai historique de la Tératologie Végétale,’
Strasburg, 186.

XX1V INTRODUCTION.

ulustrations from the immense number of recorded
facts, the principle followed has been to choose those
which seemed either intrinsically the most important,
or those which are recorded with the most care. In
addition to these public sources of information, the
author has availed himself of every opportunity that
has offered itself of examining cases of unusual
conformation in plants. For many such opportunities
the author has to thank his friends and correspondents.
Nor has he less reason to be grateful for the suggestions
that they have made, and the information they have
supplied. In particular the writer is desirous of ac-
knowledging his obligations to the Society, under whose
auspices this work is published, and to Mr. 8. J.
Salter, to whom the book in some degree owes its
origin.

The drawings, where not otherwise stated, have
been executed either from the author’s own rough
sketches, or from the actual specimens, by Mr. E. M.
Wiliams. A large number of woodcuts have also
been kindly placed at the disposal of the author by the
proprietors of the ‘ Gardeners’ Chronicle.’?

As it 1s impossible to frame any but a purely arbi-

‘ In some instances diagrams and formule are given in explanation
of the conformation of monstrous flowers; in general these require no
further explanation than is given in the text, unless it be to state that
the horizontal line — is intended to indicate the cohesion of the parts
over which it is placed, while the vertical line | signifies the adhesion
of the organs by whose side it is placed. The formula

st st ST sT st

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

INTRODUCTION, " xxv

trary definition of teratology or to trace the limits
between variation and malformation, it may suffice to say
that vegetable teratology comprises the history of the
irregularities of growth and development in plants, and
of the causes producing them. These irregularities
differ from variations mainly in their wider deviation
from the customary structure, in their more frequent
and more obvious dependence on external causes rather
than on inherent tendency, in their more sudden ap-
pearance, and lastly in their smaller lability 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.’ Following Goethe, these changes in the
course of development -are sometimes called metamor-
phoses. In this way Agardh’ 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

' 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.
* Agardh, “ Theoria Syst. Plant.,” p. xxiii.

XXV1 INTRODUCTION.

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 vertebree, &e.
3. Collateral metamorphoses, comprising those permu-
tations of form and function manifested m 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-

stances.
In collateral metamorphosis, in the same way, we

have an illustration of the degree of change possible in
ageregates of organisms under existing circumstances.

INTRODUCTION. XXVl

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
pecuhar 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. Itis 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
ageregate of organs, or of an aggregate of individuals
of whatever degree, unless we know approximately, at
least, what are the limits of each. It is not possible
to trace these limits accurately in the case of natural
science, but the larger our knowledge and the wider
our generalisations, the closer will be our approach
to the truth.

The most satisfactory classification of malformations
would be one founded upon the nature of the causes
inducing the several changes. Thus, in all organised
beings, there is a process of growth, mere increase in
bulk as it were, and a process of evolution or metamor-

XXVIII INTRODUCTION.

phosis, 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
erowth 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 :—Ilst. Arrest of Growth; 2ndly.
Excessive Growth; 38rdly. Arrest of Development ;
Athly, of Excessive or Irregular Development.

In practice, however, there are so many objections
to this plan that it has not been found practicable to
carry it out. The inability arises to a great extent
from our ignorance of what should be attributed to
arrest of growth, what to excess of development, and
so on. Moreover, a student with a malformed plant
before him must necessarily ascertain in what way it is
malformed before he can understand how it became so,
and for this purpose any scheme that will enable him
readily to detect the kind of monstrosity he is examin-
ing, even though it be confessedly artificial and im-
perfect will be better than a more philosophical
arrangement which circumstances prevent him from
employing.

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

INTRODUCTION, XXIX

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.’ It is hoped
that both the lists of books and of plants are sufficiently
full for all general purposes.”

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
attached.

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, Kirschleger, Cramer, Hallier, and
others, malformations are arranged primarily according to the organs
affected, an arrangement which has only convenience to justify it. It is
hoped that the index and the headings to the paragraphs in the present
volume will suit the convenience of the reader as well as if the more
artificial plan just alluded to had heen adopted.

? 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 ina
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.

XXX INTRODUCTION.

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 stance, to the sections treating on prolifi-
cation, and afterwards to those on multiplication. To
facilitate such research, numerous cross references are
supplied.

In the investigation of teratological phenomena con-
stant reference must be made to the normal condition,
and wice versd, 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 m determinate position
and numbers, and of regular proportionate size.
A very close approach to such a flower occurs

INTRODUCTION, XXX]

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
ordinate 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

XXX1l INTRODUCTION.

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
eroups 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.’

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 Nature.

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. Nandin in the ‘Comptes Rendus,’
1867, t. 64, pp. 929—933.

INTRODUCTION. XXXll

given individual or species ; it carries us back some
stages further im 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
flowers.

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. Sufticient has
c

XXX1V INTRODUCTION.

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

Itis probable that many terms and expressions calculated to mislead

INTRODUCTION. XXXV

That teratology may serve the purposes of syste-
matic botany to a greater extent than might at first
be supposed becomes obvious from a consideration
of such facts as are mentioned under the head of
Peloria, while the presence of rudimentary organs, or
the occasional appearance of additional parts, or other
changes, may, and often do, afford a clue to the 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
allotropism.

The mutations and perversions of form, associated
as they commonly are with corresponding changes of
function, show the connection between teratology and
physiology—a connection which is seen to be the
more intimate when viewed in the light afforded by
the writings and experiments of Gertner, 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 true
condition of affairs be thoroughly well ascertained by organogenetic
research or other means.

XXXVl INTRODUCTION.

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 lability 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 imstance, as alterations in the period
of flowering, in the duration of the several organs, and
so forth.t 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
organisation.

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

! 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 frill. Whether this be a constant
occurrence in the particular variety is not known.

INTRODUCTION. XXXVI

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 m which they
are held entirely to those peculiarities which, by an
unhappy application of words, are called monstrous
by botanists. Grafting, layering, the “ strikmg”’ 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 mherent tendency
which some plants possess of exchanging an_her-
maphrodite for a unisexual condition.

For reasons then of direct practical utility, no

d

XXXVI INTRODUCTION.

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-
tribute.

VEGETABLE TERATOLOGY.

moO ha)

DEVIATIONS FROM THE ORDINARY ARRANGEMENT
OF ORGANS.

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, m the first instance at least, the rootlets
or fibrils are arranged in regular order one over another,
in acertain 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 1s soon
lost as the plant grows.

In the case of the leaves there are two principal

l
"

2 ARRANGEMENT OF ORGANS.

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 litle
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 :
8 9 10
3 + 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 whor]
of eight. In such a case it will generally be found

hed,
Do oN

ARRANGEMENT OF ORGANS. 5

that the whorls alternate one with another, as may be
represented by this symbol :

i) 10 11 12 13 14 $615 16
Ii 2 3 A, oD 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 :

aN

i 2

Thus, while in both the annular and the spiral modes of
development the individual members of each complete
series necessarily alternate one with another, in the
former case the series themselves alternate, while in
the successive arrangement they are placed directly
one over the other. There are, of course, exceptions,
but the rule is as has been stated, and the effect is to
prevent one leaf from interferig 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-

A, ARRANGEMENT OF ORGANS.

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
orowth 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

Ce ee of five or more distinct sepals, equal

\\ In size, and arranged in a whorl, a

(Cp o \ ‘corolla of a nibs number of petals
28, ) | alternating with the sepals, five
wy, y stamens shaw: in the same position
with reference to the petals, and

SS five carpels alternating with the

Fic. 1. — Diagram stamens. Throughout this book
showing the arrange- ; :
ment of parts inacom- this arrangement is taken as the
ale leant ; a les standard of reference. Neverthe-
p, petals; st, stamens; less the spiral order does occur in
2 aaa 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;

5 ee

ARRANGEMENT OF ORGANS. 5
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, &e.

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

6 ARRANGEMENT OF ORGANS.

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 heht rather of a scaffolding,
which enables us to see the buildme 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, owimg to the
non-development of the internodes, or to excessive
erowth, or development ; but in many utstances it is

ARRANGEMENT OF ORGANS. Z

impossible, without studying the development of the
malformed flower, to ascertain whether the altered
arrangement is due to an excessive or to a diminished
action. Practically, however, it is of comparatively
httle importance to know whether, say, the isolation
of parts, that are usually combined together, is con-
genital (7.e. the result of an arrest of growth preventing
their union), or whether it be due to a separation of
parts primitively undivided; the effect remains the
same, though the cause may have been very different.

The principal alterations to be mentioned under this
head may therefore be conveniently arranged under
the following categories :—Union, Independence, Dis-
placement, Prolification, Heterotaxy, and Heterogamy.

PAAR Dials
UNION OF ORGANS.

THe union of parts, usually separate in their adult
condition, 1s 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 1s 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 imtegrity or subse-
quent coalescence of organs exist under ordinary cir-
cumstances. Considered as a teratological pheno-
menon, union admits of beimg grouped into several
subdivisions, such as Cohesion, Adhesion, Synanthy,
Syncarpy, Synophty, &c. Hach of these sub-divisions
will be separately treated, but it may be here said that,
in all or any case, the degree of fusion may be very
shght, or it may be so perfect that there may be a
complete amalgamation of two or more parts, while to
all outward appearance the organ may be single. The

COHESION. 9

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 imduce the
coalition of organs primarily distinct.

CHA PPE RR? IT:
COHESION.

Fottowrne 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
eamosepalous calyx, or of the petals in a gamopetalous
corolla. It may arise either from a union between
organs originally distinct, or more frequently from a
want of separation between parts, which under general
circumstances become divided during their 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 whereim the
union takes place between the branches of the same
plant, or between the margins of the same leaf-organ,
or between those of different members of the same
whorl.

Cohesion between the axes of the same plant—This cohesion
may occur in various manners. Firstly. The branches
of the main stem may become united one to the other.
Secondly. Two or more stems become joined together.

10 COHESION.

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

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

Fic. 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 ateazle (Dipsacus sylvestris). Union of the
branches may be the result of an original cohesion of the
buds, while in other cases the fusion does not take place
until after development has proceeded to some extent.
Of this latter kind illustrations are common where the
branches are in close approximation ; if the bark be
removed by friction the two surfaces are very likely to
become united (natural grafting). Such a union of
the branches is very common in the ivy, the elder, the
beech, and other plants. It may take place in various

FASCIATION. 1]

directions, lengthwise, obliquely, or transversely, ac-
cording to circumstances. ‘This mode of union belongs,
perhaps, rather to the domain of pathology than of tera-
tology. Some of the instances that have been recorded
of very large trees, such as the chestnut of Mount
Aitna, are really cases where fusion has taken place
between several of the branches, or suckers, thrown out
from the same original stem.’ The same process of
grafting occurs sometimes in the roots, as in Taxus
baceata mentioned by Moquin, and also in the aerial
roots of many of the tropical climbing plants, such as
Clusia rosea, &e.

Fasciation—In the preceding instances of union
between the branches, &c., the actual number of the
fused parts is not increased; but if it happen that an
unusual number of buds be formed in close apposition,
so that they are hable to be compressed during their
growth, union is very likely to take place, the more so

Fie. 3.—Fasciation in Lettuce.

from the softness of the young 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 Pinus.
‘ Regel. Garten Flora,’ vol. 8, tab. 268.

12 COHESION.

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

FASCIATION. 13

than on the other, and hence arises that curvature of
the fasciated branch so commonly met with, ¢. g.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 frequeritly in the branches ; thus, while in

Fic. 5.—Fasciated branch of Pinus Pinaster.

14 COHESION.

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 may be deformed like the parent
branch, or they may resume the ordinary aspect of the
twigs.

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

|,
Va \) $y) J fpf
N Wy,

Wl) |

“Ak f/

WAY YP

Wi yf
V

g Mad

Y

/

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

PASCIATION. 15

crest along the apex. When, as often happens, the
deformity 1s accompanied with a twisting of the branch
spirally, the buds may be placed irregularly, or in other
cases along the free edge of the spiral curve. In a
specimen of Bupleurum fale atwm mentioned by Moquin
the spiral arrangement ‘of the leaves was replaced by
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 arranged
on an ellipticalaxis. A case of this nature is described
by Schlechtendal (‘ Bot. Zeit.,’ 1857, p. 880), in Cytisus
nigricans, and M. Moquin-Tandon describes an instance
in the vine in one flower of which sepals, petals, stamens,
and ovary were abortive, while the receptacle was hyper-
trophied and fasciated, and bore on its surface a few
adventitious buds." The pedicels of Sti eptocarpus Few
have also been observed in a fasciated state.”

It has been occasionally observed that the fasciated
condition is hereditary; thus, Moquin relates that
some seeds of a fasciated Cirsiwm reproduced the same
condition in the seedlings,’ while a similar tendency
is inherited in the case of the cockscomb (Celosia).

With reference to the nature of the deformity in
question there is a difference of opimion ; 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 stemor branch. Linnzeus, 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 :—‘‘Hasciata dict solet planta cum plures caules
connascuntur, ut unus ex plurimis instar fascice evadat

1 «Bull. Soc. Bot. France,’ 1860, p. 881.

? Thid., 1861, p. 708.
3 Thid., 1860, p. 925.

16 COHESION.

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 1s
depicted in the fasciated condition.

Thestriz, which these stems almostinvariably present,
exhibit the lies 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 together. Prof. Hincks' cites cases of this kind

2 WS)
(yeas
y SpVRN)S
Ys
(

N

==

——S-
|

(f

Zs

A ) Wan
Ley) eee
ZS ( }

Dy)

iy)

Fia. 7.—Fasciation in the scape of the Dandelion (Leontodon Taraxacum),

' «Proce. Linn. Soc.,’ April 5, 1853.

FASCIATION. 17

in Primula vulgaris, Mieracitum aureum, and Ranunculus
bulbosus. I have myself met with several cases of the
kind in Primula veris, in the Polyanthus, in the Daisy,
and in the Leontodon Taravacum, 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, 1s 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 1s 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
eanal. 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

18 COHESION.

instances of the union of two, and of only two, stems
where the internal appearance was the same as in other
fasciations.

Moquin, moreover, raises the objection that it is
unlikely that several branches should become united
lengthwise in one plane only, and, further, that in the
oveater 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 DeCandolle’
of a stem of Spartiwm junceum haying several branches
only imperfectly fasciated.

Fasciated stems, then, seem to be best oxplenieal 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. Sedwm cristatum,

1 «Organ. Végét.,’ pl. iii, fig. 1.

FASCIATION,. 19

Celosia, &e. This condition may be induced by the
art of the gardener—* Lit idem arte, si plures caules
enascentes cogantur penetrare coarctatum spatiwm et par-
twrivi tanquam ea .angusto utero, sic sepe in Rainunculo,
Beta, Asparago, Hesperide Pinu, Celosid, Tragopogone,
Scorzonera Cotula fatida,” Linnaeus 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
orowth, as lettuce, asparagus, endive, &c., all of which
are very subject to this change. In the ‘ Transactions
of the Horticultural Society of London,’ vol. iv, p. 321,
Mr. Knight gives an account of the cultivation of the
cockscomb, so as to ensure the production of the very
large flower-stalks for which this plant is admired.
The principal points in the culture were the 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.

20 COHESION.

EXOGENS.

a. Herbaceous.

pe ear Oe tripartitus.
bulbosus !
Philonotis.

Delphinium elatum.

7 sp.!

Hesperis matronalis.

*Cheiranthus Cheiri!

*Matthiola incana!

*Brassica oleracea! var. pl. inflor.

Linum usitatissimum !
Althea rosea!
Lavatera trimestris.
Geranii sp.

Tropeolum majus!

Viola odorata inflor. !

Reseda cdorata!

Fragaria vesca.

Ervum lens.

Trifolium resupinatum.
repens !
pratense !

Saxifraga mutata.
irrigua.

Bupleurum falcatum.

Bunium flexuosum.

*Sedum reflexum !

cristatum !

Epilobium augustifolium !

Momordica Elaterium !

Gaura biennis.

Cotula feetida.

Barkhausia taraxacifolia.

Carlina vulgaris !

Aparyia autumnalis.

*Leontodon Taraxacum infor. !

Centaurea Scabiosa.

*Cichorium Intybus!

Hieracium Pilosella.

aureum.
umbellatum.

*Chrysanthemum Leucanthemum.

indicum !

Anthemis nobilis.

arvensis.
Cirsium lanceolatum.
Conyza squarrosa!
Inula dysenterica !
Tragopogon porrifolium.
Cnicus palustris.
Carduus arvensis !
Helianthus tuberosus !

annuus.

Cineraria palustris.
Helianthus sp. !
Dahlia variabilis.
Bellis perennis inflor. !
Coreopsis sp. !
Crepis virens.
Lactuca sativa!
Zinnia elegans.
*Campanula medium!
rapunculoides.
thyrsoidea.
Dipsacus pilosus.
fullonum.
silvestris.
Knautia arvensis.
Phyteuma orbiculare.
Jasione montana.
*Linaria purpurea !
Antirrhinum majus!
Veronica amethystea.
Veronica maritima.
sp.
Russellia juncea!
Digitalis purpurea !
Ajuga pyramidalis.
Hyssopus officinalis.
Dracocephalum moldavicum.
Myosotis scorpioides.
Echium pyrenaicum.
simplex.
Stapelie sp.
Lysimachia vulgaris!
Androsace maxima.
Primula veris inflor. !
denticulata inflor. !
Polemonium cceruleum.
Conyolvulus sepium !
arvensis !
Plantago media.
*Kuphorbia Characias.

exigua.
* Cyparissias.
Suceda maritima.

*Celosia sp.
Beta vulgaris inflor. !
Phytolacca sp.

B. Woody.

Berberis vulgaris.
Hibiscus syriacus !
Acer pseudo-platanus !
Dodonecea viscosa.
Sterculia platanifolia.
Euonymus japonicus!
Vitis vinifera inflor. !

Spartium Scoparium !
Spartium junceum !
Cytisus Laburnum.
nigricans.
Chorozema ilicifolium.
Amorpha sp.
Phaseolus sp.
Prunus sylvestris.
Laurocerasus !
Rosa sp. !
Spirea sp. !
Cotoneaster microphylla!
Ailanthus glandulosus.
*Fraxinus Ornus !
* excelsior !
Melia Azedarach.
Xanthoxylum sp. !
Sambucus nigra !
Aucuba japonica.
Erica sp. cult.
Jasminum nudiflorum !
officinale !
Olea europea.
Punica Granatum.
Tex aquifolium !

PASCIATION. i!

Daphne indica.
Daphne odora.
Suzeda fruticosa.
Ulmus campestris.
Alnus incana.
Salix vitellina, &e. !
Thuja orientalis.
Pinus pinaster !
sylvestris !
Abies excelsa !
Taxus baccata.
Larix europea.

ENDOGENS.

Lilium Martagon.
candidum !

*Fritillaria imperialis !

Asparagus officinalis !

Hyacinthus orientalis !

Tamus communis!

Narcissi sp. !

Gladiolus sp.

Zea Mays.

Filices.

See also—Moquin-Tandon, ‘Elem. Ter. Veget.,’ p. 146; C. O. Weber,
‘Verhandl. Nat. Hist.,’ Vereins, f. d. Preuss., Rheinl. und Westphal.,
1860, p. 347, tab. vii; Hallier, ‘Phytopathol.,’ p. 128; Boehmer, ‘De
plantis Fasciatis,’ Wittenb., 1752.

Cohesion of foliar organs ——This takes place in several
ways, and in very various degrees ; the simplest case is
that characterised by the cohesion of the margins of
the same organ, as in the condition called perfoliate in
descriptive works, and which is due either to a cohesion
of the margins of the basal lobes of the leaf, or to the
development of the leaf in a sheathing or tubular
manner. As an abnormal occurrence, I have met with
this perfoliation in a leaf of Goodenia ovata. The
condition in question is often loosely confounded
with connation, or the union of two leaves by their
bases. In other cases the union takes place between
the margins of two or more leaves.

Cohesion of margins of single organs —The leaves of Hazels
may often be found with their margins coherent at the

22, COHESION

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 7’%/ia, 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. Tiliv,’ Berlin, 1861.) Itis also stated
that this condition is not perpetuated by grafting.

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

Fic. 8.—Pitcher-shaped leaf of Pelargoniwm.

of the leaf is funnel-like, and the petiole is cylindrical,
not compressed, and grooved on the upper surface, as
is usually the case. A comparison of the leaves of
Pelargonium peltatum with those of P. cucullatum (‘Cavy.

OF MARGINS OF LEAVES, ETC. 25

Diss.,’ tab., 106) will show how easy the passage is
from a peltate to a tubular leaf. In these cases the
tubular form may rather be due to dilatation than to
cohesion. M. Kickx' 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* notices something of the same kind in
the leaf of Amorpha fruticosa; Treviranus® in that of
Aristolochia Sipho.

M. Puel* describes a leaf of Polygonatum multiflorum,
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, T'rifoliwm repens, where on each side of the base
of the petiole the stipules had the form just indicated.
That the bracts also may assume this condition, may be
inferred from the peculiar horn-like structures of Mar-
graavia, Which appear to origimate from the union of
the margins of the reflected leaf.

Tubular petals occur normally in some flowers, as
Helleborus, Hpimediwm, Viola, &c., and as an excep-
tional occurrence I have seen them in Ranunculus repens,
while in Hranthis hyemalis transitions may frequently
be seen between the flat outer segments of the perianth
and the tubular petals. To Dr. Sankey, of Sandywell
Park, I am indebted for the flower of a Pelargonium,

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

2 * Linnea,’ tom. 13, p. 383.

: , Verhandl. Nat. Hist. Vereins,’ 1859, Bonn, tom. xvi, tab. 3.
* «Bull. Soc. Bot. Fr.,’ vol. i, p. 62.

DA COHESION

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

Fie. 9.—LHranthis 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 m 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.’ 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 Sole anne 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

' * Report of Internat. Bot. Congress,’ London, 1866, p. 131, tab.

vii, figs. l1O—13.,
2 «Bull. Acad. Roy. Belg.,’ t. xviii, 2nd part, p. 179.

OF SEVERAL ORGANS. Fe

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 1 is the result
of fusion.

Cohesion of several organs by their margins :—leaves, &e.—'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 T’ropceolwiv
majus, and Strelitzia regina ; nm 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, ovyphylla. M. Clos’ has ob-
served the same thing in the leaves of the lentil Hrvwin
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.”

Cohesion of the leaves frequently accompanies the
union of the branches and fasciation as might have
been anticipated. Moquin cites the fenestrated leaves
of Dracontium pertusum, as well as some cases of a
similar kind that are occasionally met with, as instances

' D. C., ‘Organ. Végét.,’ pl. xvii, fig. 3, and pl. xlviii, fig. 2

2 «Mém. Acad. Toulouse,’ 1862.
> Bonnet, ‘ Recherches Us. feuill.” pl. xxi, fig. 2.

26 COHESION

of the cohesion of the margins at the base and apex
of the leaf, which thus appears perforated. This ap-
pearance, however, 1s 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 apphed.

Fusion of the edges of the cotyledons also occasion-
ally takes place, as in Hbenus cretica.t It has also
been observed in Tithonia, and is of constant occur-
rence in the seed leaves of some Mesembryanthema.
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 Crassula arborescens, observed by C.
Morren.

Fic. 10.—Two-leaved pitcher of Crassula arborescens, after C. Morren.

The stipules may also be fused together in different
' De Candolle, ‘ Mém. Lég.,’ pl. v, fig. 14.

OF THE SEPALS, ETC. 27

ways; their edges sometimes cohere between the leaf
and the stem, and thus form a solitary intra-axillary
stipule. At other times they become united in such a
manner as to produce a single notched stipule opposite
to the leaf. Again, in other cases, they are so united
on each side of the stem, that in place of four there
seem only to exist two, common to the two leaves as
in the Hop.

To the Rey. M. J. Berkeley I am indebted for speci-
mens of a curious pitcher-lke 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 retainmg its shortened form, while,
in others, the two stipules had become connate into
a cup, and all trace of the midrib was lost. The cup in
question would thus seem to have been formed from the
connation of two stipules which are ordinarily abortive.

Cohesion of the bracts by their edges, so as to form
a tubular involucre, or by their surfaces, so as to form
a cupule, is not of uncommon occurrence, under natural
conditions, and may be met with in plants which
ordinarily do not exhibit this appearance.

Cohesion of the sepals in a normally polypetalous calyx
renders the latter gamosepalous, and is not of uncommon
occurrence, to a partial extent, though rarely met
with complete. I have observed a junction of the
sepals to be one of the commonest malformations
among Orchids, indeed such a state of things occurs
normally in Masdevallia Cypripedium, &c. An illus-
tration of this occurrence is given by Mr. J. T.
Mogeridge in Ophrys insectifera, im ‘Seemann’s Journal

28 COHESION

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—Linnzus mentions the occur-
rence of cohesion of the petals in Saponaria.* 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

Olematis viticella being changed into a monopetalous

Fic. 11.—Gamopetalous flower of Papaver bracteatum.

irregular one, like the corolla of Labiates, is recorded
by Jaeger.” There is in cultivation a variety of
Papaver bracteatwm, in which the petals are united by
their margins so as to form a large cup. Under

| «Phil. Bot.,’ § 125.
2 «Nov. Act. Acad. Nat. Cur.,’ 14, p. 642, t. xxxvii.

OF THE STAMENS, ETC. 99

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 shght. 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, 7. ¢. to the formation from one original staminal
tubercle of numerous secondary ones, so that the
process is rather one of over development than of
fusion or of disjunction. ‘These conditions may be met
with as accidental occurrences in plants or in flowers,
not usually showing this arrangement. Thus, for
instance, Professor Andersson, of Stockholm, describes
a monstrosity of Salix calyculata, m which the stamens
were so united together as to form a tube open at the
top like a follicle." This is an exaggerated degree of
that fusion which exists normally in Salia 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.

30 COHESION.

Further details relating to cohesion of the various parts of the flower
are cited in Moquin-Tandon, ‘ El. Ter. Veg.,’ p. 248; ‘ Weber. Verhandl.
Nat. Hist. Verems f. d. Preuss. Rhein]. 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-hke 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,' 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 Nepenthes, &c., which occur normally
and constantly, are not here ceeds Possibly some
of the cases would be more properly classed under
dilatation or excayation.

ASCIDIA.
A. Monophyllous.

1. Sarracenia-like pitchers, formed by a single leaf, the edgesvof 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) !

*Tiha europea !

Pelargonium inquinans !

Staphylea pinnata.

Amorpha fruticosa.

Pisum sativum!

Lathyrus tuberosus.

Vicia sp.

Gleditschia sp.

Ceratonia siliqua.

Trifolium repens !

Cassia marylamdica.

' «Bull. Acad. Roy. Bruxelles,’ 1838, t. v, p. 582. ‘ Bull. Acad. Roy.
Belg.,’ 1852, t. xix, part ili, p. 437.

ASOIDIA. 31

Mimosa Lophantha.
Rosa centifolia.

gallica.
Begonia sp.
Bellis perennis!
Nicotiana sp.
Goodenia ovata!
Antirrhinum majus!
Vinea rosea.
Polygonum orientale.
Aristolochia sipho ?
Codizum variegatum var. !
Spinacia oleracea.
Corylus avellana !
Polygonatum multiflorum.
Xanthosoma appendiculatum !

2. Calyptriform or hood-like pitchers, formed by the complete union
of the margins, and falling off by a transverse fissure (as in the calyx of
Escholtzia).

Tulipa Gesneriana.

B. Polyphyllous.

1. Diphyllous, formed by the union of two leaves into a single cup,
tube, or funnel, &e. ~*

Pisum sativum (stipules)!

Crassula arborescens.

Polygonatum multiflorum.

~2. Triphyllous, formed by the union of three leaves.
Paris quadrifolia var.

Besides the above varieties of ascidia formed from the union of
one or more leaves, there are others which seem to be the result of a
peculiar excrescence or hypertrophy of the leaf. Such are some of
the curious pitcher-like structures met with occasionally in the leaves
of cabbages, lettuces, Aristolochia, &. See Hypertrophy, cup-like
deformities, &c.

In addition to other publications previously mentioned, reference may
be made to the following treatises on the subject of ascidia :—Bonnet,
‘Rech. Us. Feuilles,’ p. 216, tab. xxvi, f. 1, Brassica; De Candolle,
‘Trans. Hort. Soc.,’ t. v, pl. 1, Brassiea; Id., ‘Org. Veget., I, 316;
‘Bull. Soc. Bot. Fr,’ L, p. 62, Polygonatum ; ‘ Bull. Acad. Belg.’ 1851,
p. 591, Rosa; Hoffmann, ‘Tijdschrift v. Natuur. Geschied.,’ vol. viii,
p. 318, tab. 9, Ceratonia; C. Mulder, ‘ Tijdschrift, &c.,’ vol. vi, p. 106,
tab. 5, 6, Trifolium, Mimosa, Staphylea; ‘ Molkenboer, p. 115, t. 4,
Brassica.

Bye ADHESION.

CHAPTER AL
ADHESION.

ADHESION, so called, occurs either from actual union of
originally distinct members of different whorls or from
the non-occurrence of that separation which usually
takes place between them. It is thus in some degree a
grayer deviation than cohesion, and is generally a con-
‘sequence of, or at least 1s 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 Gap leaves in the same
spiral. cycle are adherent, a detaemn teat in the axis or
a certain amount of dicloeamon 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 1s seen under ordi-
nary circumstances in the bract of the Lime tree, which
adheres to the peduncle, also in Newropeltis, while in
Hrythrochiton hypophyllanthus 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

OF LEAVES. oo

it generally forms but a part of other and more
important deviations.

Adhesion of leaves by their surfaces——'The union of leaves
by their surfaces is not of very frequent occurrence,
many of the instances cited being truly referable to
other conditions. Bonnet describes the union of two
lettuce leaves, and Turpin that of two leaves of Agave
americana, in which latter the upper surface of one
leaf was adherent to the lower surface of the leaf next
above it, and I have myself met with similar instances
in the Saliflawet and in lettuce and cabbage leaves;
other instances have been mentioned in ‘Sazifraga,
Gesnera, Sc.

In these cases, owing to the non-development of the
internodes, the nascent leaves are closely packed, and
the conditions for adhesion are favorable, but in most
of the so-called cases of adhesion of leaf to leaf by the
surface, a preferable explanation is afforded either by
an exuberant development (hypertrophy) or by chorisis
(see sections on those subjects). Thus, when a leaf of
this kind is apparently so united, that the lower surface
of one is adherent to the corresponding surface of
another, the phenomenon is probably due rather to extra
development or to fission. There is an exception to
this, however, in the case of two _ vertically-erect
leaves on opposite sides of the stem; here the two
upper or inner surfaces may become adherent, as In an
orange, where two leaves were thus united, the 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.*” The same author mentions having seen a
remarkable example of adhesion in the involucels of
Cauealis leptophylla, the bracts of which were soldered
to the outer surface of the flowers. M. Bureau* men-

* Wydler, ‘ Flora,’ 1852, p- (ad, tab. 1x.
; ‘EL. Ter. Veg.,’ p. 254.
* Bull. Soe, Bot. Fr.,’ 1857, p. 451.

34 ADHESION

tions an instance wherein the spathe of Narcissus h-
florus 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 —T he appendicularorgans
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 by Morren
as calyphyomy, kaAvé g¢vona.! Moquin cites an
instance in Geraniwm nodosum, 11 which one petal was
united by its lower surface to one of the segments of
the calyx. A similar circumstance has been observed
in Petunia violacea by Morren. Duchartre describes
an instance wherein one of the outer sepals of Cattleya
Forbesii was adherent to the labellum.’

Adhesion of the stamens to the petals is of common occur-
rence under natural circumstances. Cassini has de-
scribed a malformation of Centawrea collina, m 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, part ii, p. 335.
“ * Bull. Soc. Bot. Fr.,’ 1860, p. 25.

OF STAMENS TO PISTILS. 35

quent occurrence among Orchids. I have observed
cases of the adhesion of the seements of the perianth to
the stamen in Ophrys aranifera, Odontoglossum, sp.
&e. It is the ordimary condition in
Gongora and some other genera. |
have seen it also in Liliwm lancifolvwm.
Some forms of Crocus, 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, bemg
partially petaloid in aspect. M. de
la Vaud' speaks of a similar union in
Tigridia pavonia. Morren® describes
a malformation of J'uchsia where 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. fyg 12, Crocus.
It should be remarked that it was Adhesion of petaloid
the outer series of stamens that were st@mens to perianth.

thus fused.®

Adhesion of stamens to pistils——'The stamens also may be
united to the pistils, as in gynandrous plants. Moquin
speaks of such a case in a Scabious; M. Clos in
Verbascum australe.s 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

‘Bull. Soc. Bot. Fr.,’ 1861, p. 147.
‘Bull. Acad. Belg.,’ vol. xviii, part ii, p. 498.

1
3 See also Prillieux, ‘ Bull. Sac. Bot. Fr.,’ 1861, p. 195.
4 *Mém. Acad. Toulouse,’ 5th series, vol. ii.

36 ADHESION.

lip of the corolla of Salvia pratensis... 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 imferus”’ seems scarcely
ever to become “ calyx swperus,”

Fic.13.—Cheiranthus while, on the other hand, the
eee enor a “calye normaliter swperus”’ fre-
open carpel. 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’ cites
an instance in a vegetable marrow (Cucumis), where a
female flower had become confluent with the branch,
at whose base it was placed, and also with two or more
flowers at the upper part of the same branch, so as to
make an oblique scar running down from the apex of
the fruit to the branch.

Synanthy— Adhesion of two or more flowers takes
place in various ways; sometimes merely the stalks

' Linnea,’ vol. ii, p. 607,
? «Journal Roy. Hort. Soc.,’ new ser., vol. i, 1866, p. 200.

SYNANTHY. 37

are united together, so that we have a single peduncle,

bearing at its extr emity two flowers placed j in approxi-
mation ver y shghtly 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 Umbellifere. Union of
this kind occurs frequently m the common cornel
(Cornus), wherein one of the lower flowers becomes
adherent to one of the upper ones. In De Candolle’s
‘Organographie Végétale,’ Plates 14 and 15, are
figured cases of fusion of the flower stems of the
Hyacinth and of a Centawrea. In other cases the
union involves not only the stalk but the flowers them-
selves ; thus fusion of the flowers 1s a common accom-
paniment of fasciation, as was the case in the Campa-
nula figured in the cut (fig. 14).

Fie. 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

38 ADHESION.

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 Pomar, Opercularia, Symphyo-
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,' in which two flowers of
a hyacinth were united together simply by means of
two seoments of the peri ianth one from each flower.
A similar occurrence has been cited by M. Gay in
Narcissus chrysanthus. In like manner the blossoms of
Fuchsias or Loniceras occasionally become adherent
merely by their surface, without mvolving any other
change in the conformation of the flowers. °M. Maugin
alludes to a case of this kind in Aristolochia 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 jomed 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 tncomparabilis | remarked a ten-parted perianth,

1 * Bull. Soc. Bot. Fr.,’ 1861, p. 159.
* Thid., 1859, p. 467.

SYNANTHY. 39

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
Napellus, figured by A. de Chamisso, ‘ Linnea,’ 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 Fic. 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

4.0 ADHESION,

three separate flowers were to be found, with the
exception of the spur belonging to the labellum of the
middle flower (figs. 15, 16).

One of the most common malformations in the
Foxglove (Digitalis) results from the fusion of several
of the terminal flowers into one. In these cases the
number of parts is very variable in different instances ;
the sepals are more or less blended together, and the
corollas as well as the stamens are usually free and
distinct, the latter often of equal length, so that the
blossom, although truly complex, is, as to its external
form, less irregular than under natural circumstances.
The centre of these flowers is occupied by a two to
five-celled pistil, between the carpels of which, not un-
frequently, the stem of the plant projects, bearmg on
its sides bracts and rudimentary flowers. (See Proli-

Fie. 17.—Synanthy 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

SYNANTHY. 4]

cited by Reinsch,' where two female flowers of Salix
cinerea were so united with a male one as to produce
an hermaphrodite blossom.

It follows, from what has been said, that the number
of parts that are met with im 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 inexcess. 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 m 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
features.

Sometimes the upper lips of two
flowers are fused into one, the two
lower remaining distinct. In other — yyg. 18-gynanthie
cases, the upper lip disappears flowersofCalceolaria in
altogether, while there are two Eo eae ae peed
lower lps placed opposite one single lower one.
another; of the stamens, some-

« Plora,’ 1858, p. 65, tab. ii.

é)>

42, ADHESION.

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

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 Polygonatwm anceps, in which
within a twelve-parted perianth there were twelve
stamens and two pistils, one four-celled, the other two-
celled; hence it would appear as if a carpel belonging
to one flower had become united to those constituting
the pistil of the adjacent one. Among Orchids this
fusion of some of the elements of different flowers,
together with the suppression of others, is carried to
such an extent as to render the real structure
dificult to decipher. Sometimes flowers of Ophrys
aranifera, at first sight seemimg 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-
eridge has observed similar phenomena in the same
species at Mentone.

Sometimes the fusion affects flowers belonging to
different branches of the same inflorescence, as in
Centranthus ruber, described by Buchenau, ‘ Flora,’ 1857,
p- 293, and even a blossom of one generation of axes
may be united with a flower belonging to another
generation. Thus M. Michalet® speaks of a case
wherein the terminal flower of Betonica alopecwros 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.

SYNANTHY. 43

as to form but a single tube. It is not uncommon,
as has been before stated, to find two corollas enclosed
within one calyx, but this is probably the only recorded
instance of the fusion of the calyx and corolla of two
different flowers belonging to two different axes.

From the preceding details, as well as from others
which it is not necessary to give in this place, it would
appear that synanthy is more lable to occur where the
flowers are naturally crowded together’ 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
erave 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
restricted.

Morren in his papers on synanthic Calceolarias,
before referred to, considers that the direction in
which fusion acts is centripetal, e.g. from the 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
case wherein the two central flowers of the capitulum of Centawrea Jacea
were united together.

4A. ADHESION.

is reversed. This remark applies particularly to imdi-
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 aurantium.
*Fuchsia var. hort. !
Cnothera sp.
Saxifraga sp.
Podalyria myrtillifolia.
Prunus Armeniaca.
spinosa.
Pyrus Malus.
Persica vulgaris.
Cratzgus monogyna.
Robinia pseudacacia.’
Gleditschia triacanthos.
Syringa persica.
Cornus sanguinea.
Viburnum sp.
*Lonicera sp. plur!
Centranthus ruber !
Valantia cruciata.
Centaurea moschata.
Jacea.
Zinnia elegans.

Zinnia revoluta.
Helianthus sp.!
Spilanthes oleracea.
Dahlia.
*Leontodon Taraxacum !
Senecio Doria.
Cichorium Intybus.
Lactuca sativa.
Anthemis retusa.
*Campanula medium !
persicifolia.
Azalea indica!
‘Vinea minor.
Atropa Belladonna.
*Solanum Lycopersicum !
*Petunia violacea !
Galeopsis ochroleuca.
Betonica alopecuros.
*Digitalis purpurea !
*Antirrhinum majus !
*Linaria purpurea!
*Pedicularis sylvatica !
*Calceolaria var. hort. !
Scrophularia nodosa.
Salpiglossis straminea.
Streptocarpus Rexii.
*Gesnera var. hort. !
Aischynanthus sp. !
Thyrsacanthus rutilans !
Anagallis collina.

SYNCARPY. A

*Primula veris ! Fritillaria imperialis !
Auricula. Agave americana.
*Primula acaulis, var. umbellata! Iris versicolor.
elatior ? sambucina.
* sinensis ! Crocus, sp.
Aristolochia Clematitis. Colchicum autumnale.
Blitum sp. Narcissus incomparabilis !
Chenopodium sp. Tazetta.
Rumex sp. biflorus.
Salix cinerea. chrysanthus.
*Hyacinthus orientalis ! *Ophrys aranifera!
Lilium bulbiferum! Calanthe vestita !
croceum, et sp. alix, pl. Oncidium bicolor.
Tulipa, sp. ornithorhyneum.
Polygonatum anceps. &e. &e,

In addition to the works before cited, additional information on this
subject may be gained from the following :—Jaeger, ‘ Missbilld.,’ p. 92.
vy. Schlechtend, ‘ Bot. Zeit.,? 1856, Robinia. Weber, ‘ Verhandl. Nat.
Hist. Vereins. Preuss. Rheinl.,’ 1849, p. 290, Primula. Hincks, ‘ Rep.
Brit. Assoc. Newcastle,’ 1838, Salpiglossis. Clos, ‘Mém. Acad. Toulouse,’
vol. vi, 1862, Anagallis. Wigand, ‘Flora,’ 1856, tab. 8, Pedicularis.
Henfrey, ‘ Botan. Gazette,’ 1, p. 280, Reseda. P. Reinsch, ‘ Flora,’ 1860,
tab. 7, Petasites. Weber, ‘ Verhandl. Nat. Hist. Vereins. f. d. Preuss.
Rheinl. u. Westphal.,’ 1860, p. 352, tabs. 6 et 7, Prunus, Persica, Campa-
nula, Tarawacum, Savifraga, Silene, Hyacinthus, &. Miquel, ‘ Linnea,’
xi, p. 423, Colchicwm. Michel, ‘'Traité 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 Caprifoliacee, Rubiacee, &c. &e.
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

46 ADHESION.

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, he 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 pee 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 iui, se. 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 acompound grape. Seringe has remarked sometimes
two, sometimes three, fruits of Ranunculus tripartitus
soldered together. He has also seen three melons
similarly joined.’ 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.” Poiteau and
Turpin have figured and described in their treatise
on fruit trees, under the name of Néfle de Correa,
four or five medlars, joined together and surmounted
by all the persistent leaflets of the calyces.’

1 «Bull. Bot.,’ tab. iii, figs. 4—6.

2 «Mém. greffe Ann. Science Nat.,’ ser. i, t. xxiv, p. 334.

3 “ Mespilus portentosa.” Poit. et Turp., ‘ Pomol. Franc.,’ liv. xxxi, p.
202, pl. 202.

SYNCARPY. 47

A very remarkable example of Syncarpy has been re-
corded by E. Koenig in which nine strawberries were
borne on one stem (Fragaria botryformis),' and a similar
malformation has been observed in the Pineapple.

When two fruits are united together they may be
of about equal size, while in other cases one of the
two is much smaller than the other. This was the
case in two cucumbers given to me by Mr. James
Salter. These were united together along their whole
length excepting at the very tips; the upper one of the
two was much larger than the lower, and contained three
cells, the lower fruit was one-celled by suppression.
Both fruits were curved, the curvature being evidently
due to the more rapid growth of the upper as compared
with the lower one.

In many of these cases, where the fruits are united

Fic. 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.

48 ADHESION.

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 Synearpy 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 m form from
what is customary, and a similar thing happens with
the tomato. In the case of stone-fruits it sometimes
happens, not only that the outer portions are adherent,
but that the stones are so likewise.

M. Roeper has observed two apples grafted together,
one of which had its stalk broken, and seemed eyi-
dently borne and nourished by the other apple ;* 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.”

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

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

1 De Cand., ‘ Phys. Végét.,’ tom. ii, p. 781.
2 Sched. de monstr. plant, ‘ Act. Helv.,’ tab. i, fig. 8.
3 «Mém. greffe,’ loc. cit., tab. xxiv, p. 334.

SYNCARPY. 49

Leg.,’ pl. 2, fig. 6; pl. 3, fig. 2.) At times these
carpels become fused together, and it becomes difficult,
when the traces of the flower have disappeared, to
ascertain whether these carpels were formed in one
flower, or whether they were the result of the fusion of
several blossoms. I have seen an instance of this kind
in a plum in which there were two carpels in the same
flower, the one being partially fused to the other. The
nature of such cases may usually be determined by an
inspection of the peduncle which shows no traces of
fusion. (See chapter on Multiplication.)

When, however, the fruits are sessile, and they
become grafted together, the kind of syncarpy is
difficult to distinguish. It may, nevertheless, be said as
a general rule that the union brought about by the
approximation of two fruits, after the fall of the floral
whorls, is never so complete or so intimate as that
determined by synanthy; and also that in those cases
where there are supernumerary carpels in the flower,
and those carpels become united together, they are
rarely so completely fused that their individuality is
lost.

An analogous phenomenon takes place not uncom-
monly in mosses, the spore capsules of which become
united together in various ways and degrees. Schimper"
cites the followmg species as subject to this anomaly :—
Buxbaumia indusiata, Leskea sericea, Hypnum lutescens,
Anomodon alternatus, Clinacium dendroides, Bryum cespi-
tittum, Brachythecium plumosum, Mniwin serratum,
Splachnum vasculosum. It has also been observed in
Trichostomum rigidulum and Hypnum triquetrum.

In addition to the authorities already mentioned, the reader may
consult Moquin-Tandon, ‘ El. Ter. Veg.,’ p. 270. Turpin, ‘Mém. greffe,
Ann. Sc. Nat.,’ ser. 1, t. xxiv, p. 334. De Candolle, ‘ Organ. Veget.,’ t. 1.
Duhamel, ‘Phys. des Arbres,’ t. i, p. 304, tab. xiii, xiv. Weber,

‘Verhandl. Nat. Hist. Vereims f. d. Preuss. Rheiml. u. Westphal.,’
1860, p. 332, tab. vi et vii.

! ‘Bull. Soc. Bot. Franc.,’ 8, pp. 73 and 351, tab. 11; and Rése, * Bot.
Zeit.,’ x, p. 410.
4.

50 ADHESION

Synspermy, or Union of the Seeds——Sceeds may be united
together in various degrees, either by their intezuments,'
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 plant.—Under this
head may be classed the union that takes place between
thestems, branches, orroots of differentplants of thesame
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
cir on waenigade 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
huge 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.”

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 respectiv ely, 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 tos this union was, hee the rede carrot

1 Nymphea lutea, AMsculus Hippocastanum, &e. See Moquin, ‘El. Ter.
Veg.,’ p. 277. 2 ©. Martins, ‘Promenade Botanique,’ p. 8.

OF AXES. 5]

(fig. 23, b), with its small overgrown part above the
junction, took the colour and large dimensions of the
white Belgian (d), which, in hke manner, with its

Fie. 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 (ed). The respective qualities of the two roots

ye ADHESION,

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. ‘Gard. Chron..,’
1849, p. 421.

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 stump{M@epends on similar
facts. In Abies pectinata, says Goeppert,’ 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.

! «Ann. Se. Nat.,’ t. xix, 1843, p. 141, tab. iv.

GRAFTING. 53

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."

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

!' «Ann. Nat. Hist.,’ ser. 2, vol. ix, tab. xvi. ‘ Phytologist,’ 1857,
p. 352, &e.

DA ADHESION.

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

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

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.’

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

1) Quoted from the ‘ Revue Hortic.’ in ‘Gard. Chron.,’ 1866, p, 386.

NATURAL GRAFTS. aT)
blue, while some produced tubers partly white and
partly blue.

Adhesion of the axes of plants belonging to different species
is a more singular occurrence than the former, and is
of some interest as connected with the operation of
grafting. As a general rule horticulturists are of
opinion, and their opinion is borne out by facts, that
the operation of grafting, to be successful, must be
practised on plants of close botanical affinity. On the
other hand, it is equally true that some plants very
closely allied cannot be propagated in this manner.
Contact between the younger growing tissues 1s 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 (Viscum), 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.’ Moquin-Tandon* relates a case
wherein, by accident, a branch of a species of Sophora
passed through the fork, made by two diverging
branches of an elder (Sambucus), growing in the
Jardin des Plantes of Toulouse. The branch of the
Sophora contracted a firm adhesion to the elder, and
what is remarkable is that, although the latter has much

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

? «El. Ter. Veg.,’ p. 289.

56 ADHESION.

softer wood than the former, yet the branch of the
harder wooded tree was flattened, as if subjected to
great pressure.': It is possible that some of the
cases similar to those spoken of by Columella, Virgil,”
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 mstances 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
erowing 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—l'his often occurs
partially in the embryo plants of the common mistleto
(Viscum), but 1s not of common occurrence in other
plants, even in such cases as the orange (Citrus), the
Cycadee, Conifere, &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
Huphorbia helioscopia, wherein two embryo plants were
completely grafted together throughout the whole length

! 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!
Pasee Daubeny, ‘Lectures on Roman Husbandry,’ p. 156.

SYNOPHTY. HY

of their axes, leaving merely the four cotyledons sepa-
rate. A similar adnation has been observed by the
same botanist in Lepidiwm sativum and Sinapis ramosa,
as well as in other plants.’ [ have met with correspond-
ing instances in Antirrhinwn majus and in Crategus
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 apples the term “ rhizo-
collesy” to this union of the roots.” 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 bemg the result of the ferti-
lisation of Huchsia coccinea (quere I’. magellanica ?) by

the pollen of F. fulgens.

1 A. P. De Candolle, ‘Organ Végét.,’ t. ii, p. 72, tab. liv, fig. 1.
2 «Bull. Acad. Belg.,’ t. xx, part 1, 1852, p. 43.

PAs i aa.
INDEPENDENCE OR SEPARATION OF ORGANS.

Unpver this head are included all those imstances
wherein organs usually entire, or more or less united,
are, or appear to be, spht 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 origmally 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 bond fide separation of parts originally united
together. In the former case, the isolation of parts
arises from arrest of development, while in the latter
it is due rather to luxuriant growth. A knowledge, as
well of the ordinary as of the unusual course, of 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 Morren' pro-
posed the general term Monosy (uovworc) for all these
cases of abnormal isolation, subdividing the group
into two, as follows—l, Adesmy (a-decuse), including
those cases where the separation is congenital ; and
2, Dialysis (d:aAvw), comprising those instances where
the isolation is truly a result of the separation of
parts previously joined together. Adesmy, moreover,

1 * Bull. Acad. Belg.,’ t. xix, part iii, 1852. p. 315.

FISSION. 59

was by the Belgian savant said to be homologous
when it occurred between members of the same whorl,
e.g. between the sepals of an ordinary monosepalous
calyx, or heterologous when the separation took place
between members of different whorls, as when the
calyx is detached from the ovary, &c. The former
case would thus be the converse of cohesion, the latter
of adhesion.

To the adoption of these words there is this great
objection, that we can but rarely, in the present state
of our knowledge, tell in which group any particular
illustration should be placed.

The terms adopted in the present work are, for the
most part, not necessarily intended to convey any idea
as to the organogenetic history of the parts affected.
Where a single organ, that is usually entire, becomes
divided the term Fission is used ; in cases where parts
of the same whorl become isolated, the word Dialysis
is employed, and in the same sense in which it is
generally used by descriptive botanists, and where the
various whorls become detached one from the other,
the occurrence is distinguished by the application of
the term Solution.

CHAPTER L.

FISSION.

WHEN an organ becomes divided it receives at the
hands of descriptive botanists the appellations cleft,
partite, or sect, according to the depth of the division ;
hence in considering the teratological instances of this
nature, the term fission has suggested itself as an 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

60 INDEPENDENCE OR SEPARATION OF ORGANS.

or ‘‘ dédoublement,” or with the “disjonctions qui
divisent les organes” of Moquin-Tandon." 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 erowing 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——T'his 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’
gives as an illustration the case of a specimen of
ftuscus aculeatus in which there occurred a division of
the foliaceous branches into two segments, reaching
as far as the insertion of the flower, but no further.
He also mentions lateral cleavage effected by a notching
of the margin, the notch being anterior to the flowers
and always directed towards their insertion. In the allied
genus Danaé, Webb, ‘ Phyt. Canar.,’ p. 320, describes
the fascicles of flowers as in ‘“‘crenulis brevibus ad
marginem ramulorum dispositis.”” Sometimes, on the
other hand, Danaé has a fascicle of flowers imserted
on the middle of the upper surface, as in fuscus.
Wigand mentions an instance in Digitalis lutea, where
the upper part of the stem was divided into six or
seven racemes; possibly this was a case of fasciation,
but such a division of the inflorescence is by no means
uncommon in the spicate species of Veronica. I have

! Loe. cit., p. 295.
? «Bull. Soc. Bot. France,’ 1857, p. 758.

FISSION, 61

also seen it in Plantago lanceolata, Reseda luteola,
Campanula medium, Epacris impressa, and a bifurcation
of the axis of the spikelet within the outer glumes in
Lolium perenne’ and Anthoranthum odoratum. In the
Kew Museum is preserved a cone of Abies excelsa,?
dividing into two divisions, each bearing bracts and
scales. A similar thing frequently occurs in the male
catkins of Cedrus Libani (fig. 25).

Fig. 25.—Bifurcated male inflorescence, Cedrus Libani.

This subdivision of axial organs is not unfrequently
the result of some injury or * mutilation, thus Duval
Jouve alludes to the frequency with which branched
stems are produced in the various species of Hquisetwm,
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 Salisburia 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

! Masters, ‘ Jourl. Linn. Soc.,’ vol. vii, p. 121.

* Cramer, ‘ Bildungsabweichungen,’ p. 4, tab. vi, fig. 4, figures a case
of the same kind in Pinus Cembra,

62 INDEPENDENCE OR SEPARATION OF ORGANS.

Mereurialis in which the leaves were deeply slashed. In
Chenopodium (Quinoa the leaves were so numerous and
the clefts so deep, that the species was hardly recognis-
able, while on a branch of Rhus Cotinus observed by De
Candolle the lobes were so narrow and so fine as to
give the plant the aspect of an Uimbellifer. Wigand
(‘ Flora,’ 1856, p. 706) speaks of the leaves of Dipsacus
fullonum with bi-partite leaves ; Moquin mentions the
occurrence of a leaf of an oleander bi-lobed at the
summit, so as to give the appearance of a fusion of two
leaves. Stemheil has recorded an instance in Scabiosa
atropurpurea in which one of the stem leaves presented
the following peculiarities. It was simple below, but.
divided above into two equal lobes, provided each with
a median nerve.’ Steinheil has also recorded a Ceras-
tiwm 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, Ulmuis -

Fic. 26.—Bifureated leaf of Lamiwm album, &ce.

campestris, and also in the common nettle, Urtica dioica,

1 « Ann. des Science Nat.,’ 2nd series, t. iv, p. 147, tab. v, figs. 3 and 4.-

FISSION OF LEAVES. 63

the leaves of which latter thus resembled those of Urtica
biloba, which are habitually bilobed at the summit. M.
Clos' mentions an instance where the terminal leaf and
first bract of Orchis sambucina were divided ‘into two
segments. The same author also mentions the leaves
of Anemiopsis californica, which were divided in their
upper halves each into two lobes—also leaves of a lentil
springing from a fasciated stem and completely divided
. into two segments, but with only a single bud in the
axil. ‘The axillary branches in hke manner showed
traces of cleavage. Fig. 26 represents a case of this
kind in Lamium album, conjoined with suppression of the
flowers on one side of the stem. I have alsoin my her-
barium a leaf of Arwm maculatwin, with a stalk single at
the base, but dividing into two separate stalks, each bear-
ing a hastate lamina, the form of whichis 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.

Fic. 27.— Bifureated leaf of Pelargonium.

Fern fronds are particularly liable to this kind of
subdivision, and they exhibit it in almost every degree,
from a simple bifurcation of the frond to the formation
of large tufts of small lobes all formed on the same
plan by the repeated forking of the pinnules. These
may be considered as cases of hypertrophy.

Moquin-Tandon, at a meeting of the Botanical Society

' *Mém. Acad. Scien. Toulouse,’ 5th series, vol. iii.

64. INDEPENDENCE OR SEPARATION OF ORGANS.
of France (April 3rd, 1858) exhibited a leaf of Cerasus
Lauro-Cerasus divided in such a manner as to resemble
a leaf of Citrus or of Phyllarthron. In this case, there-
fore, the disunion must have taken place laterally, and

;
i
Ly;

<ede
Swi

jj i LD
LZ FMA i
Swe \ AP sa
ss V4)
as \4 fa
Sn
=|=
RSs i
SHE
S
eS

if

Fic. 28.—Bifurcated frond, Scolopendriwm vulgare.

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

A. Braun describes a singular case in a leaf of Irina

FISSION OF LEAVES. 65

glabra wherein the blade of the leaf on one side was
deeply and irregularly laciniated, the other side re-
maining entire. (Verhandl., d. 35, Naturvorschef-
versamlung, tab. 3.) Lacimiate varieties of plants are
of fr equent 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 fhiese
is subjoined. A similar fission takes place constantly
in the cotyledons of some plants, sometimes, as in
Conifere, to such an extent as to give an appearance
as if there were several cotyledons.’

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 complete. 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,
Geraniwm 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. Se. Nat.,’ 3rd series, 1848, vol. x, p. 207.
i)

66 INDEPENDENCE OR SEPARATION OF ORGANS.

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

Trollius europzus dissectus.
Chelidonium majus laciniatum !
Glaucium luteum.
Brassica oleracea !
Tilia parvifolia laciniata.
asplenifohia !
Acer platanoides laciniatum.
crispum.
Assculus Hippocastanum incisum !
asplenifolium.
Vitis vinifera apiifolia!
laciniosa.
Tlex Aquifolium !
Rhus Toxicodendrum quercifolinm.
Cotinus.
Ervum Lens.
Cytisus Laburnum quercifolium !
incisum.
Rubus fruticosus laciniatus !
Pyrus communis.
Cerasus Lauro-cerasus.
Apium graveolens !
Pimpinella magna.
Saxifraga.
Cratzgus Oxyacantha laciniata.
quercifolia !
Ribes nigrum.
Sambucus nigra laciniata !
racemosa laciniata.
Dipsacus fullonum.
Scabiosa atropurpurea !
Symphoricarpus racemosus.
Helianthus sp.!
Lonicera Periclymenum querci-
folia !
Syringa persica laciniata!
Syringa vulgaris !
Nerium Oleander!
Lamium purpureum.
album !
Salvia officinalis.

Solanum Duleamara!
Fraxinus excelsior crispa.

Veronica austriaca.

Polemonium czruleum.
Juglans regia laciniata!
heterophylla.
filicifolia.
Anemiopsis californica.
Chenopodium Quinoa.
Ulmus americana incisa.
Fagus sylvatica heterophylla !
laciniata !
aspleniifolia !
incisa.
salicifolia !
Mercurialis perennis.
Urtica dioica.
Quercus Cerris laciniata !
pubescens filicina.
Betula populifolia laciniata.
alba dalecarlica.
Alnus ineana laciniata !
glutinosa laciniata !
quercifolia.
oxyacanthifolia.
Corylus Avellana heterophylla !
laciniata !
urticifolia.
Carpinus Betulus incisa !
quercifoha.
heterophylla.
Castanea vesca heterophylla.
quercifolia.
incisa.
Populus alba acerifolia. /
palmata.
quercifolia.
balsamifera.
Orchis sambucina.
Arum maculatum.
Filices sp. pl.

See also Schlechtendal, ‘Bot. Zeit.,’ tom. xiii, p. 823. A. Braun,

For Ferns too numerous for insertion, see Moore,

loc. supra citat.
Clos, ‘Mém. Acad. Toulouse,’

Nature-Printed Ferns,’ 8yo ed., 2 vols.
1862, p. 51.

Fission of the petals, &e—The floral leaves are subject
to a similar process of cleavage to that which has just

FISSION OF PETALS, ELC. 67

been mentioned as taking place in the leaves. This,
indeed, occurs very often as a normal occurrence as in
the petals of mignonette (Jteseda), or those of Alsine
media and many other plants. Here, however, we have
only to allude to those instances in which the cleavage
occurs in flowers whose sepals or petals are usually
entire. Under this category Moquin mentions a petal
of Brassica oleracea completely split into two. Linné
im his ‘Flora Lapponica’ (pp. 145 and 164) mentions
quadrifid petals of Lychnis dioica, and much. divided
petals of Rubus arcticus. Among other plants subject
to this division of sepals or petals may be mentioned
as having come within the writer’s personal observa-
tion, Ranunculus Lingua, R. acris, Papaver sommiferum,
and others of this genus, Suponaria sp. Dianthus,
Narcissus, &c.

In some of the garden varieties of Cyclamen the
corolla looks at first sight as if double, and the plan
of the flower is oblong or elliptical, instead of circular.
In these flowers each lobe of the corolla is divided
almost to the base into two lobes, so that there appear
to be ten lobes to the corolla imstead of five, as usual.
The stamens are normal in form and number in these
flowers.

In the paroquet tulips of gardeners the segments of
the perianth are deeply and irregularly gashed, the
segments occasionally becoming rolled up and their
margins coherent so as to form little tubular spurs. I
have also noticed the segments of the perianth in Crocus
and Colchicum deeply cleft, so much so sometimes, as
to equal in this particular the stigmas. In the flowers
of a species of Oncidiwm, communicated to me by Mr.
Currey, the lip 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.

68 INDEPENDENCE OR SEPARATION OF ORGANS.

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

Fic. 29.—Flower of Oncidiwm sp. seen from the back. The lip is
divided into three unequal segments.

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.’

Moquin mentions having seen the stamens of Mat-
tuola incana and Silene conica completely divided,
each section bearing half an anther, exactly as happens
in Polygalacee. In tulips and hhes the same author
mentions division of the anther only, the filament
remaining entire, as happens naturally m many species
of Vacciniwin.

A division of the individual carpels occurs very fre-
quently when those organs become more or less leafy,
as in Trifolium repens, and other plants to be hereafter
mentioned.

The instances given in this chapter have all been
cases wherein the division or the accessory growth
has taken place in one plane only and that plane

1 Masters, ‘ Rep. Bot, Congress,’ London, 1866, p. 136, tab. 7, f. 15, 16.

DIALYSIS. 69

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.

CHAPTER ILI.

DIALYSIS.

Ts 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’ in-
cluded cases of this description under the category of
*‘ Disjonctions qui isolent les organes.”’

Dialysis, as here understood, may be the result of
an arrest of development, in consequence of which
parts that under ordinary circumstances would become
fused, do not do so; or, on the other hand, it may be
the result of an actual separation between parts 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 Loc, cit., p. 298.

70 INDEPENDENCE OR SEPARATION OF ORGANS.

Dialysis of the margins of individual foliar organs—In cases
where the leaf or leaf-lhke 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 Mranthis hyemalis, wherein the
petals (nectaries) are tubular and the sepals flat, I
have met with numerous instances of transition from
the one form to the other, as shown in fig. 9, p. 24.

It is, however, in the carpels that this separation
occurs most frequently. When these organs appear
under the guise of leaves, as they often do, their margins
are disunited, so that the carpel becomes flat or
open. ‘This happens in the strawberry (fragaria), the
columbine (Aquilegia), in Trifolium repens, Ranun-
culus Ficaria, &e.

Dialysis of the parts of the same whorl:—calyx.—T'he 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, &e. When this change
happens in the calyx we have the gamosepalous con-
dition replaced by the polysepalous one, as thus
represented :

instead of

as in a calyx of five coherent sepals.

Detachment of this kind occurs not unfrequently, as
in Primula vulgaris, Trifolium repens, &e. In Rosacee
and Pomacee this separation of the calyx is of the more
moment, as it has reference to the structure of the

‘ Masters in Seemann’s ‘ Journal of Botany,’ 1867, p. 158.

DIALYSIS OF THE COROLLA. 71

inferior ovary, as will be more fully mentioned here-
after. Here, however, a case recorded by M. J. E.
Planchon may be alluded to' wherein a quince fruit (Cy-
donia) was surmounted by five leaves, the surface of the
pome being marked by as many prominences, which
apparently ‘corresponded to the five stalks of the caly-
cine leaves. In this 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 ipa so-called
inferior ovary is partly axial partly foliar, 7. e., sepaline,
and partly carpellary in its nature.

‘Dialysis of the sepals in calyces that are usually
gamosepalous has been most frequently observed in
Rlosaceew, Pomacee, Umbellifere, less commonly in
Leguininose, also in the followmg genera :—Primula,
Symphytum, Gentiana, Campanula, &e.

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 Labiate the
upper lip of the corolla may be often met with partially

Fic. 30.—Dialysis of the sepals and petals in Correa.

1 «Bull. Soe. Bot. France,’ t. xiii, 1866, p. 234.

(a INDEPENDENCE OR SEPARATION OF ORGANS.

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

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, Dahhas, &c.,
more or less deeply divided into their component
parts.

A more complete separation occurs not unfrequently
in Campanula, Rhododendron, Phlox, Fc. 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, CU. 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 anew 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 Lhododendron. fk. linearilobum, a Japanese
species, offers a good illustration of this.

The following list contains the names of the genera

DIALYSIS OF THE CARPELS. 73

in which this separation of the petals of an ordinarily
gamopetalous flower takes place most frequently.

Correa. Verbaseum !
Campanula! sp. pl. Mimulus.
Polemonium. Digitalis !
Phlox! Orobanche.
Cobeea! Solanum.
Rhododendron ! Nicotiana.
Erica! Gentiana !
Rhodora. Anagallis.
Azalea ! Primula!
Composite! sp. pl. Lamium!
Lonicera ! Convallaria !
Convolvulus ! Lilium !
Pharbitis. Colchicum !
Antirrhinum ! &e. &e.

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
(Malvacew) become double, one of the first stages of
the process is often the disjunction of the stamens, and
a similar dissociation occurs in Leguininose and Con-
posite, as in Tragopogon, as related by Kirschleger,
in Hypochaeris by Wigand, and in Coreopsis by Schlech-
tendal.

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

Generally speaking, the disunion is complicated with
frondescence—but not always so. I have, in my her-
barium, specimens of Convallaria majalis, Commelyna
sp., and of LInliwm auratum, mm all of which the three
carpels are completely disjoined, and present three

1 ‘Bull. Bot.,’ pl. i, figs. 8—12.

74 INDEPENDENCE OR SEPARATION OF ORGANS.

styles, three stigmas, &c., without any other change.
Engelmann’ speaks of three classes of this malformation.
Ist, 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
Moguin in Stachys sylvatica, in which, owing to im-
perfect disjunction, the two bi-lobed carpels were
changed into a nearly one-celled capsule ;’ and 35rd,
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
Cerasus and Amygdalus persica; nevertheless, fleshy
fruits sometimes become disunited. I have seen a
case similar to that mentioned by M. Alphonse de
Candolle in Solanum esculentum, in which the pericarp
became ruptured, and the placentas protruded. <A
hke occurrence has also been observed in a species of
Melastoma.? This is analogous to what happens in
Caulophyllum and Slateria. Disjanction of the carpels

Fic. 32.—Anomalous form of orange.

* «De Anthol.,’ p. 37.
; Moquin, loc. cit., p. 305.
‘ Neue Denkschr. der Allg. Schweiz. Gesell.,’ band v, pl. ii, p. 5.

DIALYSIS OF THE CARPELS. 75

is not rare in oranges. Sometimes this takes place
regularly, at other times irregularly; occasionally in
such a manner as to give the appearance of a hand and
fingers to the fruit. Of one of these, Fer -ari,' in the
curious volume below cited, speaks thus: “ Arbor pro-
fusissima, quia dat utraque manu; imo quia vere manus
dat in poma conversis ; utque magis munifica sit poma
ipsa convertit In manus.”

M. Duchartre’ 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® 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.

.

Fic. 33.—Orange. Showing dis- Fic. 34.—Section of orange
junction of carpels, after Maout. 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

' * Hesperides,’ auctore Ferrario, Rome, 1646, fig. 415, pp. 213 and
215. See also Michel, ‘ Traité du Citronnier.’

? « Ann. des Science Nat.,’ 5rd series, 1844, vol. i, p. 294.

® *Org. Véget.,’ vol. ii, p. 41.

76 INDEPENDENCE OR SEPARATION OF ORGANS.

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.

CHAPTER HT.
SOLUTION.

THE isolation or separation of different whorls that
are ordinarily adherent together is by no means of rare
occurrence. Were it not that the isolation is often 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 hetérologue” of Morren.
Moquin Tandon does not make any special subdivision
for the class of cases here grouped together, but places
them all under ‘‘ Disjonctions qui isolent les organes.”
It seems, however, desirable to have a separate word
to express the converse condition of adhesion, and for
this purpose the term solution, as above stated, is here
employed. Diagrammatically, the condition may be
expressed by placing a dotted line at the side of the
letters thus :

CNG pie 6. SBS e
CG eed 0a) e)50) a
would indicate the disjunction of the sepals from the

SOLUTION OF CALYX FROM PISTIL. 77

carpels (C), in contradistinction to adhesion, which may
be represented by the unbroken lne thus:

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 adherens 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 ftosacee, 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 (fosa) 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 Pomacee. 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-

78 INDEPENDENCE OR SEPARATION OF ORGANS.

flowered thorn, Cratequs Oxyacantha, in some blossoms
of which the hollowed end of the peduncle still invests

Fra. 35.—Proliferous Rose. Showing an absence of the usual 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."

This condition accords precisely with the account

1 «Gard. Chron.,’ 1865, p. 554; 1867, p. 599.

SOLUTION OF CALYX FROM PISTIL. 79

of the development of the flowers in Pomacee as given
by Payer, Caspary, and others, so that the flowers

Fie. 36.—Section through Apple Fie. 37.—Calyx detached from
blossom, showing detachment of carpels in Apple.
ealyx from ovaries, absence of
dilated flower-stalk, &e.

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 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
leaves.

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.

80 INDEPENDENCE OR SEPARATION OF ORGANS.

normal organogeny as well as from unusual 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 Rosacee are concerned, there is so
much variation in the relative position of calyx and
carpels under ordinary circumstances, that it 1s no
matter for surprise that similar diversities should exist
in teratological cases. A similar remark will apply to
Savifragacee, Cucurbitacee, Myrtacee, Bruniacee, Rubi-
acece, and other families of like conformation.

In Umbelliferw, 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-

Fie. 38.—Flower ition of the fruit. |
of @nanthe crocata, : : ;
eatin” Pag In the Rubiaceous genus Bikkhia,
sepals were com- ag mentioned by Duchartre, the ovary
Pees pees, is completely inferior, but when the
here three in num- fruit arrives at maturity four small
Pacteaet °F leaves are detached from its surface

which had previously adhered to it,
and which it seems reasonable to consider as the
sepals.

In Campanulacee a similar separation of calyx from
the ovary may be occasionally met with. On the other
hand, the occasional formation of a leaf on the inferior
ovary of these plants would indicate the axial nature
of the fruit. In Campanumea and Cyclodon the calyx
is inferior, while the corolla is superior. In the last-
named genus this peculiarity “is carried to the highest
degree, the sepals being, in C. parviflorwm, placed on
the peduncle of the flower far removed from the base
of the corolla and ovary, whilst in C. truncatwm and in

SOLUTION OF CALYX FROM PISTIL. 8]

Campanumea they adhere to the base of the tube of the
corolla.”! In this order, then, as in Savifragacea,
Bruniacee, &c., no hasty seeiusion should be drawn
as to the nature of the fruit. In Brunia microphylla
the oyary is superior, enclosed within but not adnate
to the cup-like calyx, to which latter, however, the
petals and stamens are attached.

In Onagracee (Jussieua), as also in Cactacee (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 Kpi-
lobium, I have met with four perfect leaves at the
summit of the ovary, in the place usually occupied
by the sepals. This ‘would also favour the notion that
the axis entered into the constitution of the fruit in
this genus.

Mr. B. Clarke, in his ‘ New Arrangement of Phanero-
gamous Plants,’ p. 4, cites a case wherein the perianth
was completely detached from the surface of the ovary
in Cannabis sativa.

It must be borne in mind that some of the recorded
instances of change in the relative position of the calyx
and pistil ought more properly to be referred to a sub-
stitution of carpels for stamens, as in Begonia, Fuchsia,
&e. Among Cucurbitacee, examples have been re-
corded, both of the detachment of the calyx from the
ovary, and of the partial conversion of some of the
anthers of the male flower to carpels.

The very singular mode of germination of Sechiwm
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.

'! Hook et Thoms, ‘ Precurs. ad Flor. Ind.,’ Journ. Linn. Soc., vol. 11,
1858, p. 6.
* Lindley, ‘ Veget. Kingd..’ p. 315.
6

82 INDEPENDENCE OR SEPARATION OF ORGANS.

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

*Rosa! Athamanta.
*Pyrus! Selinum.
*Crategus ! Carum.
*Daucus ! Imperatoria.
Pastinaca. Rudbeckia!
Torilis. *Campanula!
Apium. Lonicera !
Cnanthe ! Cucumis !
Heracleum, Cannabis.

Solution of the stamens from the petals——A separation of
the stamens from the petals in flowers, wherem those
organs are usually adherent one to the other does not
often occur unattended by other changes. It has been
observed in Cobea scandens (Turpin), in Antirrhinun
majus, and in many double flowers.

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

| Masters, ‘ Journal of Linnean Society,’ 1866, vol. vii, p. 207. On
the subject of this chapter the reader should also consult Moquin-
Tandon, 1. c., p. 298. Engelmann, ‘De Antholysi,’ p. 37, tab. v. C.
Morren, ‘Bull. Acad. Belg.,’ xix, part 5, 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, ‘Traité général de Botanique, p. 57.
Bentham, ‘Journ. Linn. Soc.,’ vol. x, p. 104 (Structure of Myrtacee),
and other treatises on Organography,

PART Tir.
ALTERATIONS OF POSITION.

NECESSARILY 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
Heterogamy.

4 ALTERATION OF POSITION,

CHAPTEHR=%.
DISPLACEMENT.

Rea 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 disunions, suppressions, hypertrophies, &.,
are very liable to bring about or to be accompanied
with changes in the position, either of the parts
directly affected or of adjoining organs.

In this place, then, it is merely necessary to allude
to some of the more important displacements, and to
refer for further details to the sections relating to those
irregularities of growth on which the displacement
depends.

Displacement of bulbs—I owe to the kindness of Mr.
James Salter a tulip bulb which had been dug up after
flowering, and from the base of which were suspended
several small bulbs; and I have smce 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 seales of the parent organ, and which, m some
way or another, have been displaced and thrust into the
ground. Professor de Vriese figures something of the
same kind in [via carminosa.'

Of somewhat different nature to those above described
was an anomaly described by M. Gay at a meeting
of the Botanical Society of France, April 8th, 1859.
The plant affected was Leucoium cstivum, and the
changes observed were apparently attributable to a
simple separation of two leaves that are usually con-
tiguous. “Suppose,” says M. Gay in describing this

1 «Tijdschr. voor. nat. Gesch.,’ viii, 1841, tab. ii, p. 178.

DISPLACEMENT. 85

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 evident
thickening of the upper portion of the lengthened
internode, and there will be not only a single bulb,

Fria. 39.—Unusual position of bulbs of tulip; the parent-bulb cut 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 the head of hypertrophy.

Displacements affecting the inflorescence——T'hese 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

86 ALTERATION OF POSITION.

characters used to distinguish genera ; thus one of the
distinctions between rye-grass (Lolivm) 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 Lolium
with one edge against it; but in a specimen of rye-
erass that has come under my own observation, the
arrangement was that of Triticwm.

M. Kirschleger relates having found a specimen of
Leucanthemwm pratense, m which the hgulate female
flowers were growing singly in the axils of the upper
leaves of the stem.’ 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, Dipsacee,
&c. Ihave also met with specimens of Lamium albwin
in which some of the fascicles or clusters of flowers in
place of being placed at the same level on opposite
sides of the stem were placed alternately one above
another.

Caspary’ 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 corre esponding deficiency. Another flower of
the same plant bore on its pedicel a small leaf, which
was doubtless the bract raised above its ordinary
position.

M. Fournier mentions an instance in Pelargoniwm
grandiflorum, where, owing to the lengthening of the
axis, the pedicels, instead of being umbellate, had
become racemose; and I owe to the kindness - 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 Internat. Bot. Congress, Paris, 1867.
2 «Bot. Zeit.,’ 1859, p. 117, tab. v.,

DISPLACEMENT OF LEAVES. 87
axis. There were several bracts beneath this flower
disposed spirally in the 3 arrangement, all being empty,
excepting the terminal one. In like manner, a head
of flowers becomes sometimes converted into an umbel.

Displacement of leaves—A cohesion of parts will some-
times give rise to an apparent displacement, but the
true nature of the malformation can, in general, be
readily made out.

Steinheil’ found a specimen of Salvia Verbenaca, the
leaves of which presented very curious examples of
displacement arismg 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 hmb 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.” The term “diremption” has sometimes
been applied to cases where leaves are thus apparently
drageed out of position.

1 « Ann. Sc. Nat.,’ ser. 2, vol. iv, 1835, p. 143, tab. v.
* See Kirschleger, ‘ Flora,’ 1844, p. 566 (Scabiosa).

88 ALTERATION OF POSITION. E

In Tradescantia virginica 1 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-

Fie. 40.—Large-coloured leaf oceupying the position of the inflores-
cence in Gesnera, after Morren.

DISPLACEMENT OF LEAVES. 89

trophy, whether that 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,’ where, in
Gesnera Geroltiana, a large leaf apparently occupied

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

| + Bull. Acad. Belg.” t. xvii, part 1. p. 387.

90 ALTERATION OF POSITION.

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 mstance, in some conifers, such as
the Larch, (Abies Larix) or Pinus pinea, there may be
found at different stages in the growth of the branches
leaves in crowded fascicles or tufts; while, when
erowth is more rapid, the leaves may be disposed in
a spiral or alternate manner.

In the yew (Taaus) the leaves at the ends of the
shoots not unfrequently lose their usual distichous
arrangement and become arranged in a close spiral

Fia. 42.— Altered arrangement of leaves of yew, Tawus baccata.

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

DISPLACEMENT, 9]

So, too, the alteration from’ verticillate to spiral, or
vice versd, may take place without any other notable
change.t This may frequently be seen in Rhodo-
dendrons.

Displacement of the parts of the flower—This subject 1s
partly touched on in the chapters on solution, ad-
hesion, and in those on_ hyper trophy, elongation,
prolification, &e., so that in this place it is only requi-
site to offer a few general remarks, and to refer to
other sections for further details. Morren, in referring
to displacement of the floral organs, mentions an
instance in a Fuchsia, wherem 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.”. The same author
also gives an account of the displacement of several of
the organs of the flower in Cypripedium insigne, the
displacement being consequent, apparently, on a spiral
torsion proceeding from right to left, and involving the
complete or partial suppression of several of the organs
of the flower. The dislocation of organs im a spiral
direction led Morren to apply the term ‘‘ speiranthie ”’
to similar deviations from the usual construction.
Changes of this kind among Orchidacee are by no
means uncommon ; the following may be cited by way
of illustration. Ina specimen of Oncidinm cucullatum
furnished me by Mr. Anderson, well known for his
success as a cultivator of Hie 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
appreciated by the accompanying formula than even

' «Clos. Mem. ee Toulouse,’ 5th ser., t. vi, pp. 51, 70.
* «Bull. Acad. Roy. Belg.,’ xviii, part ii, p. 505, and vol. xvii, part 1,
p. 196, and vol. xix, part i, p. 260.

92 ALTERATION OF POSITION.

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

s standing for sepal, p for petal, 1 for lp, sr for
stamen; then the dislocated form may be represented
thus :

1E IF

Frag. 43.—Flower of Oncidium cucullatum, showing union of two lower
sepals, displacement of column and lip, &e.

In a specimen of Cypripedium also furnished by Mr.
Anderson the appearance was as represented in the
accompanying figure and diagrams, figs. 44, 40.
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

DISPLACEMENT. ' 93

complete absence of the lower one. In the second or
inner whorl of the perianth the lip is merely a little
oblique on one side, but the lateral petals are distorted,
displaced, and adherent one to the other and to the

/ I J \ \

Fic. 45.—Diagram of mal-
formed Cypripedium. 0, outer
segments; 7, Inner segments of
perianth ; e, lip; s, stigma; a
anther.

>

Fic. 44.—Malformed flower of
Cypripedium.

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

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

e
Wig amare Me ‘aay
4 = \ eae
N} \ A . ————S a

{ a (STG
“\\ CF ) “78 \ \
\ SS v4 fj eo) Ve

¥ offal (WY

\ \e 4

— WS

Fic. 46.—Diagram showing ordi-
nary arrangement in Cypripedium.
o, outer, 7, Inner segments of peri-

Fic. 47.—Plan of flower of Ly-
caste Skinneri, showing displace-
ment of organs.

anth; e, lip; a, anther; a’, abortive

stamen; s, stigma.

were present, one of them with a long tubular spur, 00 ;
of the two lateral petals, 77, 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

OA. ALTERATION OF POSITION.

column and ovary of this flower were in them normal
condition.

Cohesion of two or more segments of the perianth
is frequently associated with displacements of this
nature: thus, in a flower of Dendrobiwn 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

ate / |
(? oN a ( ")

Fie. 48.—Plan of malformed Fia. 49.—Plan of natural arrange-
flower of Dendrobium nobile. ment in Dendrobiwm 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
isP p parts in this species.
CoN Sometimes the displacement seems
if ~ ) consequent on hypertrophy of one of
Ds the parts of the flower, the dispro-
\ ‘Js portionate size of one organ pushing
> WV é
the others out of place. This was
Fig. 50.—Plan of the case in a violet, fig. 50, in which
flower of violet iN ae ] ei
showing displace. ONC 0 e sepals s was greatly
aa of petals, a thickened, and the petals and stamens
t was a rudi- rf 2 : : x
ment ofa stamen. Were displaced in consequence.
It is curious to observe in many
of these cases that the transposed organ not only

DISPLACEMENT. 95
occupies the place of a suppressed or abortive organ,
but frequently assumes its colour, and, to some extent,
its function. ‘This has been alluded to in the case of
the leaf of Gesnera (see p. 88) and in Orchids this
replacement seems to be very common; thus, in
addition to the cases before mentioned, in a flower
of an Odontoglossum, for which | am imdebted to
Professor Oliver, the two lateral sepals were united
together and occupied the position of the labellum,
which was absent. A similar occurrence happens
occasionally in Lycaste Skinneri, thus recalling the
structure of Masdevallia, where the labellum is normally
very small. ‘The arrangement in Lycaste may thus
be symbolised :

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 ; 1t seems as though, when the labellum, which
performs so important an office in attracting and guid-
ing insects, 1s deficient, its place is suppled by other
means.

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 Primula sinensis and in the
Pea (Piswin 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

96 ALTERATION OF POSITION.

central axis of the flower, after the fashion of a spiral
tube.’

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-
located.

Displacement of the placentas aud 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 bemg placed on the
suture, or rather on the margins of the altered carpel,
are placed on the surface of the expanded carpel.
Thus, in some double flowers of Ranunculus Ficaria that
came under the writer’s notice the carpels were open,
i.e. disunited at the margins, and each bore two 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.’

The occurrence, also, of different forms of placen-

' See also Schlechtendal, ‘ Bot. Zeit.,’ iv, p, 804. Primula veris, parti-
bus perigonii spire in modum confluentibus.
= Seemann’s ‘ Journal of Botany,’ vol. v, 1867, p. 158.

DISPLACEMENT. 97

tation in different flowers on the same plant is no
unusual thing in malformed flowers ; thus, in double
flowers of Saponaria officinalis I have met with sutural,
parietal, and free central placentation in the same plant.'

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”
(fig. 51). See also Lindley, ‘ Veg. Kingdom,’ p. 497.

Fie. 51.—1. Monstrous flower of a Cerastium; sepals and petals
leafy. 2. Stamens and pistils separate. 3. Ovary cut open to show the
imperfect dissepiments and the attachment of the ovules. 4. A deformed
ovule.

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

! «Journ. Linn. Soc.,’ i, 1857, p. 161. ¢. aylog.
* * Adansonia,’ ii, 306.

~

98 ALTERATION OF POSITION.

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

The same excellent observer has recorded the occur-
rence of free central placentation in malformed flowers
of Trifoliwm repens."

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 Gesneracee.

The same author alludes to certain changes in the
same flower where two open carpels ‘‘ were soldered
together laterally, as was clear by the rudiments of
two styles, the placenta being produced only at the two
united edges, the outer margims 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, im the
absence of which the spongy receptacle of the seeds
must have been supposed to spring from the medial
nerve

In other cases the placentas were parietal above, but
axile at the base of the capsule, a striking instance of
the facility with which axile placentation becomes
parietal, the change being here effected by the pro-
longation of the axis, and the formation on it of a
second whorl of carpellary leaves.

In double flowers of Primulacee 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” an instance of true foliar.

! * Adansonia,’ iv, p. 70, t. 1.
? Henfrey’s *‘ Bot. Gazette,’ 1, 265.

DISPLACEMENT OF PLACENTAS. 99

and true axile placentation in the same flower in Vinca
minor.

These and many similar changes, which it is not
necessary further to allude to, are not so much to be
wondered at when it is borne in mind how shght an
alteration suffices to produce a change in the mode of
placentation, and how frequent is the production of
adventitious buds or of folhar outgrowths, as may be
seen in the sections relating to those subjects and to
Substitutions.

It will be remembered, also, how, in certain natural
orders, under ordinary circumstances, considerable
diversity in placentation exists, according as the 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 Caryophyllee, Cucurbitacee,
Papaveracee, and many other orders, varies according
to the age of the carpel, and if any stasis or arrest of
development occurs the placentation becomes altered
accordingly.

It is not necessary, in this place, to enter into the
question whether the placenta is, in all cases 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 fohar 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 :

RB. . Brown, ‘ Ann. Nat. Hist.,’ 1843, vol. xi, 35. Brongniart, ‘Ann. Sc.
Nat.,’ 1834, sér. 2, i, p. 308. Alph. De Candolle, ‘Neue Denkschrift der

100 ALTERATION OF POSITION.

Allg. Schweizer Gesellsch.,’ Band v, 1841, p. 9. Duchartre, ‘Ann. Se.
Nat.,’ 3rd ser., 1844, vol. ii,p. 290. Ibid., ‘Elem. Bot.,’ p. 574; ‘ Rev.
Bot.,’ 1846-7, p. 213. Babington, ‘ Gard. Chron.,’ 1844, p.557. Lindley,
‘Elements,’ p. 89; ‘Veg. King.,’ pp. 313, 497, &e. Berkeley, ‘ Gard.
Chron.,’ 1850, p. 612. Unger, ‘Nov. Act. Acad. Nat. Cur., 1850;
and in Henfrey’s, ‘Bot. Gazette,’ 1851, p. 70. Schleiden, ‘Prin-
ciples,’ English edit., p. 385. Payer, ‘Elem. Bot., pp. 196, 211,
224. Baillon, ‘Adansonia,’ ili, p. 310, tab. iv. Cramer, ‘ Bildungsab-
weichungen,’ p. 20, &e. Clos, ‘ Ann. Se. Nat.,’ 5th ser., ii, 313, as well
as any of the general treatises on botany. Reference may also be made
to the chapters on Prolification and Substitutions (in the case of the
carpels and ovules), and to the authorities therein cited.

CHART BR ET:
PROLIFICATION.

Moaqutn-Tanpon and other writers have classed the
production of buds in unwonted situations under the
head of multiplication, but, as the altered arrangement
is of graver import than the mere increase in number,
it seems preferable to place these cases under this
heading rather than under that of alterations of
number.

The adventitious bud may be a leaf-bud or a flower-
bud ; it may occupy the centre of a flower, thus 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-

PROLIFICATION. 10]

mation than the median form. If only the number of
orders and genera be reckoned, the truth of this state-
ment will be scarcely recognised ; but if individual cases
could be estimated, the ditference in frequency between
the two would be very much more obvious. ‘This may,
perhaps, be explained by the fact that the branch has
a greater tendency to grow in length than it has to
develop buds from the axils of the leaves. The flower
is admitted to be homologous with the branch, and it is
also known that, up to a certain time, the branch-bud
or leaf-bud and the flower-bud do not essentially differ."
At a later stage the difference between the two 1s
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 hmiting the length of the central
axial portion. Now, in prolified flowers the functions
and, to a considerable extent, the appearance of a leaf-
bud or of a branch are assumed, and with them the
tendency to grow in length is developed. Median pro-
hfication, therefore, in this sense, is a further step in
retrograde metamorphosis than is the axillary form.
To grow in length, and to produce axillary buds, are
alike attributes of the branch; but the former is much
more frequently called into play than the latter; for
the same reason, median prolification is more common
than the axillary form. This is borne out by the
frequency with which apostasis, or the separation of
the floral whorls one from another, to a greater degree
than usual, is met with in prolified flowers.

In both forms the adventitious growth is much more
frequently a flower-bud or an inflorescence than a leaf-
bud or a branch. This may be due to the position of
the flowers on a portion of the stem of the plant espe-
cially devoted to the formation of flower-buds, to the
more or less complete exclusion of leaf-buds, 7. 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

' Linn., ‘ Prolepsis,’ § vii; Goethe, ‘ Metamorph.,’ §$ 96, 103, 106.

102 ALTERATION OF POSITION.

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
hable 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, &e.
From the similar arrest of growth in leneth, 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
inflorescence.

Median prolification occurs frequently in double
flowers; the axillary variety, on the other hand, 1s 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
isa useful points of comparison, and may therefore
be passed over,

Prolification of the inflorescence.—'l'lis consists in the for-
mation of leaf-buds or of an undue number of flower-

PROLIFICATION OF THE INFLORESCENCE. 103

buds on the inflorescence. It must be distinguished
from virescence, or the mere green colour of the floral
organs, and from chloranthy, in which all or the greater
portion of the parts of the flower are replaced by leaves.
Prolification is, in fact, a formation of supernumerary
buds, leafy or floral, as the case may be, these buds
being sessile or stalked, the ordinary buds beige not
necessarily changed. Prolification of the inflorescence,
like the other varieties, admits of subdivision, not only
according to the fohar 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 lght of
a malformation’ seeing that a similar condition is so
commonly metwith normally, as in Epacris, Metrosideros,
Bromelia, Eucomis, &c., wherein the leafy axis projects
beyond the inflorescence proper; or as in Primula im-
perialis, m which plant, as also in luxuriant forms of
P. sinensis, tier after tier of flowers are placed in succes-
sion above the primary umbel. Nevertheless, when we
meet with such conditions in plants which, under
ordinary circumstances, do not manifest them, we must
consider them as coming under the domain of teratology.

Median foliar prolification of the inflorescerce is frequently
met with in Conifere, 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 Cryptomeria
Japonica, has enabled me to verify nearly the whole of
Caspary’s observations. A similar prolongation of the
axis occurred in some of the male catkins of Castanea

* “ Diaphysis inflorescentiarum,”” Engelmann, ‘ De Anthol.,’ § 85.

104 ALTERATION OF POSITION.

vesca, each of which had a tuft of small leaves at their
extremity. In the common marigold and in Lotus
corniculatus I have also seen instances of this kind.
Kirschleger' describes a tuft of leaves as occurring on
the apex of the flowering spike after the maturation of
the fruit in Plantago, and a similar growth frequently
takes place in the common wallflower, in Antirrhinwm
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, im which the
spike was surmounted by a tuft of leaves and roots, as
well as a still more singular instance in Hryngiwm vin-
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 Mélicoq’ gives a case in
Primula variabilis, in which at the top of the flower-
stalk, in the centre of six flowers, was placed a complete
plant in miniature, having three leaves, from the axil
of one of which proceeded a rudimentary flower. Mr.
W. B. Jeffries also forwarded me a polyanthus (fig. 52)
in which the peduncle was surmounted by a small plant,
forming a crown above the ordinary flower-stalk, Just -
as the crown of the pineapple surmounts that fruit. A
similar instance was exhibited at the Scientific Com-
mittee of the Horticultural Society on July 11th, 1868,
by Mr. Wilson Saunders; the species in this case was
P. cortusoides. To Mr. R. Dean I am indebted for a
similar proliferous cyclamen, which seems similar to
one mentioned by Schlechtendal.* This author alludes
to an analogous circumstance in the ‘inflorescence of:
Cytisus nigricans, where, however, the change was not
‘Flora,’ 1844, p. 565.

1
2 «Ann. Se. Nat.,’ ser. 3, vol. v, 1846, p. 64,
* * Bot. Zeit.,’ vol. xx, p. 382.

PROLIFICATION OF THE INFLORESCENCE. 105

so great as in the preceding cases. The instances just
cited all occur in plants having an indefinite form of

Fie. 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. “ fi
raceni,’ says he, “ultra flores producuntur in ramos,
foliosos duo hifidos qui tandem trium wnciarum longi-
tudinem adepti fuerunt.’*

. Median floral prolification of the inflorescence, wherein
a new inflorescence projects beyond the primary
one, is not uncommon in plants having thei flowers
arranged in close heads or umbels; as in the common

' * Miscel. Austriac. Bot.,’ vol. i, Vindob, 1778, p. 133.

106 ALTERATION OF POSITION.

wild celery and other Umbellifere.". 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 Tvriticwm repens, Roéper m Hu-
phorbia palustris.’

Lateral foliar prolification of the inflorescence 18 of more com-
mon occurrence than the preceding. I have met with
it, amongst other plants, frequently in Brassica oleracea,
Pelargonium zonale, Scabiosa, Bellis, and many other
composites, also in Leguminose, e.g. Lupinus, Trifolium,
Ooronilla, &c. Prof. Oliver forwarded me a specimen
of Huphorbia geniculata in which, m 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 im 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 Serophularia 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. Many species of Alliwm, Lilium, Saxifraga,
Begonia, Achimenes, normally produce leaf-buds or bulbs
in the inflorescence ; so, too, leafy shoots are sometimes

1 “Umbellati dum prolificantur, augent wmbellulam, ut ex umbellula
simplici altera eweat.” ‘Linn. Phil. Bot.,’ § 124.

2< En. Kuphorb.,’ p. 36.

3 Meisner, ‘Mon. Gen. Polygoni Prodrom.,’ p. 20, tab. v, considers the
bulbils of this plant to be modifications of the pedicels of the flower.

PROLIFICATION OF THE INFLORESCENCE, 107

found in Alisma natans, Juncus uliginosus, Chlorophytum
Sternbergianum, &c. As an accidental occurrence, a
similar thing has been noticed in Lychnis coronaria,
Phaius grandifolius, Oncidiwm cebolleta, EHpidendrwm
elongatum,' &e. &e.

Here, too, 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 re originate in this situation, and in either
case the inference is that the ovary is, in part at least,
made of the dilated and hollowed axis. Leaves may
occasionally be found in this way on the so-called
calyx-tube or on the inferior ovaries of roses, pears,
apples, Pereskia, Crategus tanacetifolia, &e.

The fruits of Opuntia Salmania and of O. fragilis
(‘ Bull. Soc. Bot. France,’ vol. i, p. 306; vol. v, p. 115)
have been observed to form small fruit-like branches
around their summits. This circumstance is more fully
treated of in the succeeding chapter relating to Hete-
rotaxy.

Lateral floral prolification of the infloreseence.—T'his, which is
termed by Hngelmann Hcblastesis foliorum sub flora-
lium,” is much the most common of all these deviations,
and it is met with in every degree, from the presence of
a single supernumerary flower in the axil of a bract
to the existence of a small cluster or panicle of such
flowers.

Itis common inthe Anemone coronaria and hortensis,
also in the common scarlet Pelargonium (fig. 53). It has
been frequently recorded in Poteriwm sanguisorba, and
in Sanguisorba officinalis, and 1s especially common in
Umbellifere, Dipsacece, and Composite ; a familar 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.

2 “ Prolificatio e latere ex calyci communi proles plurimos pedunculatos
emittens, fit in compositis aggregatis proprie dictis.” ‘Linn. Phil. Bot.,’
§ 124,

108 ALTERATION OF POSITION.

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

Fria. 53.—Lateral prolification in inflorescence of Pelurgonium.

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 Sepiwm 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' gives a summary of the various kinds of malfor-
mation affecting the inflorescence in Plantugo, and

divides them into five groups, as follows:—1st, bracteate,
wherein the inferior bracts are quite leaf-like, as is
frequently seen in Plantago major. 2nd, roseate ; bracts
leafy in tufts or rosettes, without flowers, as in the
so-called rose plantain, common in old-fashioned
gardens in this country. 3rd, polystachyate; spike-
branched, bearing other spikes in the axils of the bracts,
as in P. lanceolata, P. maritima, &c. 4th, proliferous,
where the flower-stalk bears a rosette, a spike, or a

1 «Bot. Zeit.,’ 1857, p. 873. See also ‘Verhandl. Nat. Hist. Vereins.
Preuss. Rheinl. u. Westphal.,’ 1854, t. 1x.

PROLIFICATION OF THE INFLORESCENCE. 109

head with other rosettes, 5th, paniculate, in which the
inflorescence has become a much-branched pyramidal
panicle, covered with little bracts, and with very rudi-
mentary flowers.’ 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 witha branched spike provided with
large leafy bracts, the branches of the
spike being but little less in diameter
than the ordinary single spike. These
specimens would therefore seem to
be intermediate between Schlechtendal’s
bracteate and polystachyate divisions.
Wigand* 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 d
Braun would fall under the roseate sec- Fra. 54.—Plan-
tion, as would also that of Kirschleger, 92, mayor, with
panicled inflo-
though we are expressly told that the rescence.
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 haye had the opportunity of

1“ Pannicula spicatim sparsa onusta imnumera fetura herbaceorum
flosculorum racematim coherentium,” ‘ 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.

110 ALTERATION OF POSITION.

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

Fia. 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 mternodes 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. /anceolata

PROLIFICATION OF THE INFLORESCENCE. 11]
mentioned by Dr. Johnston,’ wherein were several
spikes, some sessile, others stalked and pendent, the
whole intermixed with leaves and disposed in a rose-
like manner. I have myself gathered specimens of
this nature, occurring in the same plant, at Shanklin,
Isle of Wight (fig. 56).

Fic. 56.—Inflorescence of Plantago lanceolata, bearing a tuft of leaves
and flowers at the end of the flower-scape.

It is rather singular that each species of Plantago
seems to have its own perverse mode of growth; for
instance, the bracteate, polystachyate and paniculate
forms are almost exclusively confined to P. major, the
roseate form to P. media, the proliferous form to
P. lanceolata.

The instances wherein flower-buds originate from the
surface of an inferior ovary, as in those cases where the
top of the stem is dilated so as to form part of the
fruit, would be properly classed under the head of pro-

' * Flora of Berwick-on-Tweed,’ vol. i. p. 38.

ig ALTERATION OF POSITION.

lication 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 feseda luteola (fig. 57), might equally
well be placed with fission or multiplication of the

Za
Ye.

= S20
lk

=>.
~

F1a@. 57..-Branched inflorescence of Reseda luteola.

axile organs. Branched spikes of this character are
not so common among Orchids as might be expected.

PROLIFICATION OF THE INFLORESCENCRH. 113

Professor Reichenbach enumerates a few instances in
the Report of the International Botanical Congress of
London, 1866, p. 121, and the same author gives an
illustration in his ‘ Or chidographia Europea, tab. 150.

In Grasses, as indeed in other plants with a spicate
inflorescence, this change occurs not unfrequently.
The common Ray Grass “(Lolium) i 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:

Leafy. _ Floral.
Ranunculacex. Ranunculus. Ranunculus!
Anemone. Anemone.
Crucifere. *Brassica !
Caryophyllacee. Lychnis !
Dianthus !
Geraniacee. *Pelargonium ! *Pelargonium !
Leguminos. *Trifolum ! Trifolium !
Lotus! Lotus!
Coronilla !
Cytisus. Cytisus.
Rosacee. Poterium.
*Pyrus! *Pyrus !
*Crategus ! Crategus !
*Rosa. Rosa!
Sanguisorba.
Philadelphacez. Philadelphus.
Crassulaces. Sempervivuns.
Echeveria.
Crassula.
Ficoidez. ? Tetragonia.
Cactaceex. Opuntia. Opuntia.
Pereskia.
Saxifragacer. Saxifraga !
Umbelliferse. Seseli.
*Apium !
Cnidium.
Cet ae.
Eryngium. ryngium.
2 Stans
Heracleum ! Heracleum !
Hydrocotyle. Hydrocotyle.
Daucus.

8

114

‘ Begoniacee.
Valerianacee.
Dipsacee.

Composite.

Campanulacez.
Lobeliacee.
Hricacez.
Convolvulacez.

Scrophulariacee.

Gesneracee.
Primulacez.

Plumbaginacee.
Plantaginacee.
Polygonacez.
Euphorbiacee.
Urticacez.
Amentacez.

Conifer.

Orchidacez.

Liliacee.

Leafy.

Cnanthe.
Begonia !
Valeriana.
*Scabiosa !
Knautia !

Calendula.

Tragopogon !

Bidens !

Prismatocarpus.

Azalea !
Convolvulus !

Scrophularia !
Antirrhinum !
Achimenes!
Primula !
Cyclamen !

*Plantago !
Polygonum !
Euphorbia !

Castanea!
*Larix !
*Cryptomeria !
Taxodium !
Phaius !
Epidendrum !
Oncidium !

* Allium !
*Ornithogalum !

ALTERATION OF POSITION.

Floral.

Carum.
Selinum.
Angelica !
Conium.
Astrantia.
(nanthe.

*Scabiosa !
Knautia!
*Bellis!
Centaurea.
Calendula.
Anthemis.
Coreopsis.
Apargia.
Lampsana.
Carlina.
Arnoseris.
Tragopogon !
Rudbeckia !
Senecio !
Carlina.
Pyrethrum.
Filago.
Hedypnois.
Cirsium.
Lactuea.

Jasione.

Convolvulus!
Calystegia !

Primula!
Cyclamen !
Armeria.
*Plantago !

Ficus.
Corylus !
Castanea.

Pinus.
Ophrys!

PROLIFICATION OF

FLOWER. 115

Leafy. Floral.
*TLilium !
Amaryllidacez. Fourcroya
Alismacez. Alisma !
Palmacezx. Cocos.
Juncaceez. *Juncus!
Restiacee. Restio! Restio !
Elegia ! Elegia !
Willdenovia ! Willdenovia !
Cyperacee. Carex.
Graminacee. Dactylis.
*Lolium !
Festuca.
*Zea !
*Triticum !
*Hordeum !
Secale.
Phleum.

In addition to the papers already cited the following
works may be consulted with reference to prolification
of the inflorescence :

Moquin-Tandon, ‘ El. Ter. Veg.,’ p. 376. Engelmann, ‘ De Antholysi,’
§ 85—87. Fleischer, ‘Missbild. Versch. Cultur. Pflanz.’ For figures
of Hen and Chicken Daisy (Bellis prolifera), see Lobel, ‘ Ic.,’477. Sweert,
‘ Florileg.,’ pl. 98, f.5. ‘Hort. Eystett. Plant. Vern.,’ fol. iv, f. i, &e.
For similar malformations in marigold (Calendula), see Lobel, ‘ Ic.,’ 553.
‘Act. Acad. Nat. Cur.,’ vol. x, p. 208. Jaeger, ‘ Missbilld.,? 192-195.
‘Hort. Eystett.,’ pl. estiv. fol. ui, fi. Klinsmann, ‘ Linnea,’ 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, ¢, P. major rosea, bracteata
paniculata, prolifera, &c. ‘ Hort. Hystett.,’ pl. estiv., t. vii, f. 2, P.
rosea et P. bracteata. Lobel, ‘ Stirp. Hist.,’ p. 162. Dodonzus, ‘ Pempt.,’
1—4, cap. xxiii, P. major spica multiplex, i.e. paniculata. Gerard,
‘Herbal.’ Clusius, ‘Plant. Rar. Hist.,’ lib. v, p. 109—10, Plantago
augustifolia Gareti prolifera. Marchand, ‘ Adansonia,’ iv, p. 156.
Conifere.—Richard, ‘Mem. Conif.,’ tab. xiii, f. 9. A. Braun, ‘ Das
Individ.,” 1853, p. 65. De Cand., ‘Organogr.,’ tab. xxxvi. Wigand,
* Bot. Untersuch.,’ 154. Schlechtendal, ‘ Bot. Zeit.,’ 1859, p. 239. Caspary,
‘De Abiet. flor. fem. struct. morphol.’ Parlatore, ‘Ann. Sc. Nat.,’
1862, vol. xvi, p. 215. Cramer, ‘ Bildungsabweich.,’ p. 4, &c., &e.
Graminee.—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,

116 ALTERATION OF POSITION.

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- Fic. 59.—Flower of Verbascum
berry prolonged into a leafy with five disunited sepals, five
branch. From the ‘ American similar green petals, and a pro-
Agriculturist.’ 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 obseryation.
In this case the petals were virescent, and the stamens
and pistils were entirely absent, hence in truth, the so-

PROLIFICATION OF THE FLOWER. 117

called flower more nearly resembled a branch. In a
flower of a May Duke cherry, for which I am indebted
to Mr. Salter, there was a gradual change from the
floral to the foliar condition; thus there were five dis-
tinct lanceolate sepals, the arrangement of whose
veins betokened that they were leaf-sheaths rather
than perfect leaves, ten petals partly foliaceous and
sheath-like as to their venation, one of them funnel-
shaped, but whether from dilatation or cohesion of the
margins could not be determined. The stamens were
eight or ten in number, their connectives prolonged
into foliaceous or petaloid appendages, so that the
filament represented the stalk of the leaf. The pistil was
entirely absent and its place was supplied by a branch
with numerous perfectly formed stipulate leaves.

Some flowers of Anagallis arvensis described by Dr.
Marchand’ 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.

! * Adansonia,’ vol. iv, 1864, p. 150, tab. vii. ‘Gard. Chron.,’
November 19th, 1864.

118 ALTERATION OF POSITION.

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, bearimg at
its tip two whorls of leaflets, alternately three and
three.

5. In this case two out of the five stamens were
normal, the other three changed into leaves, showing
clearly the origin of the leaflets, in the last case, which
took the place of the stamens.

6. The ovary varied in different flowers. In some
the placenta was crowned with ovules; in others the
ovules were replaced by a single whorl of leaflets; m
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 giving out a number of adventitious roots at its

base.”
In other genera of the same order (Primulacec) an

PROLIFICATION OF THE FLOWER. 119

extension of the placenta into a leafy branch has been
observed, as in Lysimachia, where in one case the pro-
longed placenta was removed and struck as a cutting.'

In Ericacee too, the axile placenta has been seen
ovuliferous 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 arepresentation of a Ranunculus affected with me-
dian floral prolification.* 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 sidé, and one of its lobes had
become petaloid. Between the calyx and the corolla
were three or four spathulate, hairy, bract-like organs ;
the corolla and stamens were unchanged ; but in place
of the usual four-lobed ovary there was a single carpel
with a basilar style, terminated by a forked stigma.
Occupying the place of the other lobes of the pistil
was an oblong woolly flower-bud, consisting of calyx,
corolla, and stamens, but with no trace of pistil. I

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

2 Baillon, ‘ Adansonia,’ i, 286.

° See also figure in ‘ Hort. Eystett. Ic. Plant. Vern.,’ fol. 15, fig. 1.
Ranunculus asiaticus.

120 ALTERATION OF POSITION.

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

gogo)

Fig. 60.— Flower of Dianthus 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 (Hpacridacee) 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 origimated 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 followimg orders seem to be the
most frequently affected by it: Ranwnculacee, Caryo-

PROLIFICATION OF THE FLOWER. 121

phyllacew, Rosacee ; while it is commonly met with in
Scrophulariacee, Primulacee, and Umbelliferw. Of genera
which seem peculiarly lable to it may be mentioned
the following: Anemone, Ranunculus, Cheiranthus,
Dianthus, Dictamnus, Daucus, fosa, Geum, Pyrus,
ni Antirrhinum, Digitalis, Primula.

A reference to the subjoined list of genera affected by
this malformation, and the knowledge of itscomparatively
greater frequency in some than in others of them, will
show that 1t is more often met with in plants having an
indefinite form of inflorescence than in those having a
definite one. The change may affect some only, or
the whole of the flowers constituting an inflorescence ;
and though it is by no means a constant occurrence, it
very frequently happens that the central or terminal
flower in a definite imflorescence is alone affected, the
others remaining in their ordinary condition, as in
pinks (Dianthus); and in the indefinite forms of inflo-
rescence, it 1s equally common that the uppermost flower
or flowers are the most lable 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 Primulacee 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 in Thesiwm also, another genus with

’ Duchartre, ‘Ann. des sc. nat.,’ 3me série, vol. ii, 1844, p. 293.

£22 ALTERATION OF POSITION.

free central placenta, this malformation has been
found.

So also among plants with indefinite inflorescence,
prolification seems very frequently to affect those
wherein the axis is normally prolonged; thus it is
common in Dictamnus, which plant has an internode
supporting the pistil; it is frequent among Umbelli-
fere, where the carpophore may be truly considered an
axile production; it 1s common among /tosacew and
Ranunculacee, 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 Magnoliacee, 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 Gewm rivale (a plant very lable 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 Leguminose and in Santalacee. In
Leguminose, as also in Anvygdalus, it would seem as if

PROLIFICATION OF THE FLOWER. 123

the adventitious bud were strictly a lateral and axillary
production, and moreover that the carpel itself is not
strictly terminal but lateral in position, though 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 Thesiwm, the pistil
was altogether absent, and occupying its place was
the new bud or branch.’

Fie. 61.—Daucus Carota, showing leafly carpels, prolification, &e.

' Reissek, ‘ Linnea,’ vol. xvii, 1843, p. 641, tab. xix.

124 ALTERATION OF POSITION.

As the carpels are not unfrequently absent in cases
of median proliication, it has been thought that the
pistil in such cases was metamorphosed into a stem
bearing leaves or flowers. 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 m 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 Daucus Carota gathered in Switzer-
land (fig. 61), not only was the calyx partially detached
from the pistil, but the carpels themselves were leaf-
like, disjomed, 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.’

' The tube of the calyx in these specimens was traversed by ten ribs,
apparently corresponding to the primary ridges of the normal fruit;
these ribs were destitute of spines, and the bristly secondary ridges
were entirely absent. Those portions of the carpels which were detached
from the calyx had each three ribs, a central and two lateral ones, which
appeared to be continuous with the ribs of the calyx below,—although in
the case of the calyx there were ten, in the case of the carpels six ribs,
three to each. This diversity in number is thus explained :—A circle of
vascular tissue ran round the interior of the calyx-tube, at its junction
with the limb, and at the point of insertion of the petals and stamens.
The vascular circle seemed to be formed from the confluence of the ten
ribs from below. Of the five ribs in each half of the calyx, the three
central ones were joined together just at the point of confluence with the
vascular circle, above which they formed but a single rib—that traversing
the centre of the carpellary leaf; the two lateral ribs of each half of the
calyx seemed to be continuous, above the vascular rim, with the lateral
ribs of the carpel; these lateral ribs were connected on either side with

the central one by short branches of communication. The disposition
of the ten ribs may be thus represented :—

1 1 i 1 1 1
323 2 3 32323
1 Me ee PF Pad

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

PROLIFICATION OF THE FLOWER. 125

Not only are the carpels thus frequently separated
one from the other by the prolonged axis, but they
undergo commonly a still further change in becoming
more or less completely foliaceous, as in the Daucus
just mentioned, where the carpels were prolonged into
two lance- shaped leaves, whose margins in some cases
were shehtly 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 Anchusa ochroleuca, 1 which
the pistil consisted of two leaves, situated antero-pos-
teriorly on a long internode, ae a small terminal
flower-bud between them ; and numerous similar in-
stances might be cited.

In this ‘place may also be noticed those imstances
wherein the placenta elongates so much that the peri-
carp becomes ruptured to allow of the protrusion of
the placenta, altnough this prolongation is not attended
by the formation of new buds. Cases of this kind
occurring in Melastoma and Solanwm have been put on
record by M. Alph. de Candolle.' This is a -change
analogous with that which occurs in some species of
Leontice or Caulophyllum, as commented on by Robert
Brown. See ‘ Miscellaneous Botanical Works’ of this
author, Ray Society, vol. 1, 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 joi 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.
' «Neue Denkschriften der allgemeine Schweizerischen Gesellschaft,’

band 5, 1841, tab. 2.

126 ALTERATION OF POSITION.

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

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

N

\

Fig. 62.—Median floral prolification, &c., in flower of Delphiniwm.

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

PROLIFICATION OF THE FLOWER. 127

assumed from the disunion of their margins somewhat
of the appearance of leaves, other flowering branches
proceed—axillary prolification. If, on the other hand,
the carpels be few in number, and placed in a verticillate
manner, the axis then generally passes upwards without
any change in the form or position of the carpels being
apparent, as in a proliferous columbine, figured in the
‘Linnean Transactions,’ vol. xxiii, tab. 34, fig. 5.
When a flower with the ovary naturally inferior or
adherent to the calyx becomes prolified, a change in
the relative position of the calyx and ovary almost
necessarily takes place, the latter becoming superior
or detached from the calyx; this has been already
alluded to in Umbellifere. Ina species of Campanula
examined by me, the calyx was free, the corolla double,
the stamens with petaloid filaments, and in the place
of the pistil there was a bud consisting of several
series of green bracts, arranged in threes, and 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 Campanula, the styles were present, forming below
an imperfect tube which surrounded the adventitious
bud; in another, contrary to what occurs usually in
such cases, the ovary was present in its usual position,
but surmounted by a bud of leafy scales, enclosed
within the base of a tube formed by the union of the
styles. A similar relative change in the position of
the calyx and the ovary takes place when the 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
Rudbeckia ? showing this detachment of the calyx from
the ovary. In a monstrous Fuchsia that I have had
the opportunity of recently examining, the calyx was

128 ALTERATION OF POSITION.

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 along its ovuliferous margins. Such
instances as these seem to be the first stages of a
change which, carried out more perfectly, would result
in the formation of a new bud on the extremity of the
rolonged axis.

In Or chidacee, among which family I have now met
with several instances of pr olification, the ovary seems
usually to be absent. Fig. 63 shows a prolified flower

Fic. 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 mimature raceme. This
specimen was forw arded to me by Dr. Moore, of
Glasnevin.

As might be expected, it very rarely happens that
median prolification occurs without some other cleviation
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

PROLIFICATION OF THE FLOWER. 129

interest. Fusion of two or more flowers in association
with prolification is especially common in cultivated
specimens of Digitalis purpurea; the uppermost flowers
of the raceme become fused together so as to form one
large, regular, erect, cup-shaped corolla, to the tube
of which the stamens are attached, in greater number
than ordinary, and all of equal length; the bracts and
sepals are confusedly arranged on the exterior of the
flower; while in the centre, in the place usually 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 (Dianthus) 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 1s 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

: 2

130 ALTERATION OF POSITION.

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 Gewm vrivale, where the sepals
are usually large and leaf-like, as they likewise are
frequently in proliferous roses and pears.

UG) CESS
a DD7, E /

4 “4
ley IN

* iy i
Gj f
oe

Fie. 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 FLOWER. 13]

gated (fig. 64) after the fashion of Geum rivale, &c.'
Occasionally from the middle of the outer surface of the
urn-shaped thalamus proceeds a perfect leaf, which
could hardly be produced from the united sepals or
calyx-tube ; a similar occurrence in a pear is figured in
Keith’s ‘ Physiological Botany,’ plate ix, fig. 12.

The change which the calyx undergoes when flowers
with an habitually adherent ovary become prolified, and
wherein the calyx is disjoined from the ovary, has been
before mentioned, but it may also be stated that, under
such circumstances, the constituent sepals are frequently
separated one from the other, and not rarely assume
more or less of the appearance of leaves, as in pro-
lferous flowers of Umbellifere, Campanulacee, Com-
posite, &e.

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 s, deemed the doubling to be
an almost necessary precursor of prolification ;’ but,
though frequently so,"it 1s not invariably the case that
the flower so affected is double—e. g. Gewm. 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 Cheiran-
thus? mm 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

' Bell Salter, ‘Gard. Chron.,’ March 13th, 1847, and ‘ Ann. Nat. Hist..,’
1847, vol. xix, p. 471, Ke.

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

[32 ALTERATION OF POSITION.

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
Helianthemum.

‘he stamens are subject to various changes in pro-
lified 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
Umbellifere 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 heen 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 mtra-
carpellary ; the axis is so shghtly 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 Boraginee. 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

PROLIFICATION. OF THE FLOWER. 133
this form of prolification are, no doubt, those which
are met with among Primulacee and other orders with
free central placentation.

Duchartre, in his memoir on the organogeny of
plants with a free central placenta, in the ‘ Ann. des
Se. Nat.,’ 3 sér., 1844, p. 290, among other similar
instances, mentions two flowers of Cortusa Matthioli,
wherein the placenta was ovuliferous at the base; but
the upper portion, instead of simply elongating itself
into a sterile cone, had produced a little flower with
its parts slightly different from those of the normal
flowers. M. Alph. de Candolle has likewise described
somewhat similar deviations, and one in particular in
Primula Auricula, where the elongated placenta gave
off long and dilated funiculi bearing ovules, while other
funiculi were destitute of these bodies, but were much
dilated and foliaceous in appearance.’ In some flowers
of Rhododendron I have observed a similar condition
of the ovules, which, moreover, in the primary flowers,
were attached te the walls of the carpels—parietal
placentation.

In speaking of these as cases of intra-carpellary
prolification, it is, of course, impossible to overlook
the fact that they differ in degree only from those
cases where the lengthened axis projects beyond the
cavity of the carpels; nevertheless they seem to
demand special notice, because in these particular plants
the placenta or its prolongation appears never to pro-
trude beyond the carpels, or at least very rarely.
There are, however, numerous instances of such an
extension of the placenta and of prolification occurring
among Primulacee im conjunction with the more or
less complete arrest of growth of the carpels.” 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

' A. de Candolle, ‘Nene Denkschriften,’ op. cit., p. 9; also Unger as
cited in ‘ Botanical Gazette,’ May, 1351, p. 70.
* Duchartre, op. cit.

134 ALTERATION OF POSITION.

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 im the guise of small leaves.

Under the term prolification of the fruit two or
three distinct kinds of malformation appear to have
been included. The term seems usually to be apphed
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 prolification m no wise
differs from ordinary central prolification. 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.’ In other cases,
doubtless, the carpellary whorl is alone repeated, the
other whorls of the adventitious flower bemg com-
pletely absent.

Another condition, apparently sometimes mistaken
for prolification 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 prolification. This
happens frequently in oranges, as in the variety called
Mellarose.”

1 * Ann. Se. Nat.,’ 1844, vol. i, p. 297.
> Maout, ‘ Lecons Elémentaires de Botanique, vol. 11, p. 488; Ferrari,
‘ Hesperides,’ pls. 271, 315, 405,

PROLIFICATION OF THE FRUIT. 135

Moquin has given an explanation of the St. Valery
Apples, wherein the petals are sepaloid, the stamens
absent, and where there is a double row of carpels,
by supposing these peculiarities to be due to “a
prolification combined with penetration and fusion
of two or more flowers,” but it is surely more 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-hke 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.’

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-
lification, 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 Moguin-Tandon, loc. cit., p. 386, &c.; see also Trécul, in the ‘ Bull.
Soc. Bot. France,’ tom. i, p. 307.

136 ALTERATION OF POSITION.

before mentioned. Moquin, in the tenth volume of
the ‘Bulletin of the Botanical Society of France,’
p- 73, says that when the case is one of prolification
the lower fruit is larger and is formed of a fleshy
mass ; moreover, the line of demarcation between the
fruits is more distinct, and there are traces of the seed-
bearing cavity in the interior, and of calycine lobes at
the top. On the other hand, if the case be one of
hypertrophy merely, the lowermost fruit is the
smallest, and there is no trace of seed-bearing cavity
nor of sepals. See also under Hypertrophy.

Some other malformations usually referred to 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.

Ranunculacez. Clematis.
Anemone! * Anemone!
Ranunculus ! *Ranunculus !
Delphinium.
Caltha.
Aquilegia !
Crucifere. Bunias.
*Cheiranthus !
Erucago.
*Matthiola!
Sisymbrium !
Brassica !
Nasturtium.
Hesperis.
Sinapis!
Diplotaxis.
Lunaria.
Erysimum,

Cistacee.
Caryophylle.

Violacez.
Tilacez.
Geraniacee.
Sapindacee.
Malvacee.

Malpighiacez.
Rutacex.
Resedacez.

Aurantiacee.
Vitacee.
Umbellifere.

Rosacez.

? Leguminosz

Cucurbitacee.
Passifloracez.
Philadelphacez.
Onagracee.
Epacridacee.
Ericacee.

Convolvulacez.
Gentianacee.
Apocynacee.
Jasminacee.

Scrophulariacez.

Orobanchacee.

PROLIFICATLON OF

Leafy.

Dianthus !

Triumfetta !

Pavia !

Genera not specified.

Vitis

* Rosa!
*Geum !
Agrimonia.
Spirea.

*Pyrus!

Epilobium!
Epacris !

Convolvulus.

Gentiana.

Verbascum !

FLOWER,

Floral.

Alyssum.
Peltaria.
Cardamine !
Cleome.

Helianthemum !

*Dianthus !
Silene !
Lychnis !
Viola!

Geranium !
Pavia !
Paritium.
Hibiscus !
Byrsonima!
*Dictamnus!
Reseda.
Caylussa !
*Citrus !
Vitis.
Heracleum.
Angelica.
Thysselinum.
* Athamanta.
*Daucus !
*Torilis.
*Rosa !
*Geum !
Amyegdalus.
Prunus !
Spireea !
Rubus.
*Pyrus !
Trifolium !
Medicago!
Melilotus.
Pisum !
Cucumis.
Passiflora.

Philadelphus.

*Hrica.

Rhododendron !

Gentiana.
Vinca.
Jasminum !
Antirrhinum !
*Digitalis !
*Linaria !
Veronica.
Orobanche.

138 ALTERATION OF POSITION.

Leafy. Floral.
Labiate. Genera not specified. Stachys.
Phlomis !
Hydrophyllacez. Hydrophyllum.
Boraginacee. Anchusa.
Symphytum.
Primulacee. *Dodecatheon. *Cortusa.
* Anagallis ! *Anagallis !
*Primula.
Dipsacee. Scabiosa.
Composite. Hieracium! Hieracium !
Cirsium. Cirsium.
Hypocheris. Calendula !
Spilanthes.
Carthamus.
Coreopsis.
Campanulacez. Campanula. ‘“*Campanula !
Polygonacez. Genera not specified. Rumex.
Santalacee. Thesium.
Liliacee. Genera not specified. Tulipa !
Hemerocallis !
Asphodelus.
Hyacinthus !
Iridacee. Iris.
Amarylldacez. Narcissus !
Leucojum.
Orchidacee. Orchis !
Habenaria.
Cyperacee. Carex.
Graminee. Phleum.

Axillary prolification 1s 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

PROLIFICATION OF THE FLOWER. 139

interior of the flower, or from the inflorescence beneath
the flower.

The accessory bud presents itself as a leaf-bud, a
branch, a flower-bud, or a miniature inflorescence ;
it may be sessile, but is far more frequently stalked,
and in more than half the number of cases it is a flower-
bud or an inflorescence. ‘There may be one or more
of these buds; if two only, then they are usually
placed directly opposite one to the other, on the
opposite sides of the flower.

It will be seen, from the appended list, that the
orders and genera in which this description of adventi-
tious growth occurs most frequently are the following :
—Crucifere, especially the genus Brassica; Caryo-
phyllacee, e.g. Dianthus; fesedacee; Leguminose,
e.g. Melilotus, Trifolium, &c.; Rosacew, e.g. Rosa,
Potentilla, &c.; Umbellifere, and Campanulacee.
For the most part, these are groups also peculiarly
hable to central prolification.

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 aml 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-lke. 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 properly 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 mdividually,
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 im gamopetalous

14.0 ALTERATION OF POSITION.

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 hable 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
higher.

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. Caryophyllacee, Potentilla, Anemone,
Dictamnus, Umbellifere, &e. 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 disc, or else presents one or more

PROLIFICATION OF THE FLOWER. 14]

glandular swellings, e.g. feseda, Nymphea, Crucifere.
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 lable to this affection; but as
information on this point is but rarely given in the
records of these cases, | 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 1s 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 fohar 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 Caltha palustris, Herreria
parviflora, and other plants. Dr. Marchand’s flowers
of Anagallis, previously referred to at p. 117, showed
good illustrations of this occurrence, as also some
specimens described by Kirschleger in A. phenicea.'
Steinheil has figured and described’ a flower of Scabiosa

' * Bull. Soc. Bot. Fr.,’ 1863, vol. x, p. 461.

2 «Ann. Sc. Nat.,’ 1835, p- 65. See ates Le Maout, ‘ Lecons Element.,’
yol. ii, p. 426.

142 ALTERATION OF POSITION.

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, 7.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
apples to cases where leaf-buds occur on the outer
surface of inferior ovaries, as in fosacew, Pomacee,
Philadelphus, or Tetragonia expansa, as elsewhere men-
tioned.

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’ 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 m the
axils of the sepals than in the other “whorls. In
Orucifere 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. 1, p. 39, fig. 2

PROLIFICGATION OF THE FLOWER. 143

with the petals, which, together with the stamens and
pistil, remained unaffected. he number of parts in
the supernumerary structures is generally less than
the normal flowers.

In Mr. Herbert Spencer’s ‘ Principles of Biology,’
part iv, p. 37, are figured and described some mon-
strous inflorescences in Angelica and other Umbellifere,
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.*

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 Nymphea (fig. 65). The ovary in this case
was wanting, but in its place was a tuft of small leaves.
It is curious that among Dr. Kirk’s drawings of east
tropical African plants now at Kew, there should be
one representing a precisely similar state of things.
‘The species in both instances was Nymphaea Lotus, or
a cultivated variety of it.

M. Wesmael’ describes a very singular case of what
appears to have been referable to axillary prolification
in the flowers of Carex acuta. The rachillus is de-
scribed as prolonged through the utricle by the side of
the stigmas, bearing on its rside a bract, then a secon-
dary utricle, from the axil of which Spi short stem
surmounted by an ovary. Wigand, ‘Flora,’ 1856,
mentions a similar change in Carex glauca. In this
instance the base of the female inflorescence bore
lateral spikes, which projected from the utricles ; some
of these adventitious spikes were female, others

1 See also ‘ Nat. Hist. Review,’ 1865, p. 377.
? «Acad Roy. Belg.,’ April 11th, 1865.

14.4 ALTERATION OF POSITION.

female below and male above, others, again, wholly
male.
Various changes in the form and arrangement of

Fia. 65.—Flower of Nymphcea Lotus, var., showing axillary floral pro-
ification. 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-hke im appear-
ance. The dislocation of the affected sepal from its
fellows is a very frequent occurrence; in cases of this

PROLIFICATION OF THE FLOWER. 145

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

The petals, under such circumstances, often exist
in the guise of sepals or of small leaves ; and instances
are recorded wherein the place of the calyx and corolla
was supplied by a succession of overlapping green
scales, from the axils of which the new buds arose.
M. Germain de Saint Pierre records such a case in
Trifolium repens, wherein the calyx and corolla were
replaced by overlapping scales, in the axils of each one
of which arose a flower; above there was a row of
stamens, and in the centre a pistil in the guise of a
trifohiate leaf." 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 Mesa 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-
sation.

So far as the andrecium is concerned, the stamens
either remain unaltered, or they are present in a more
or less petal-like condition; but it far more frequently
happens that the stamens are entirely suppressed, the
adventitious bud supplying their place; thus was it in
the Dianthus represented in the adjoming woodcut, fig.
66, where the stamens were entirely absent, and their
places supplied by flower-bearing branches. This
Dianthus has the more interest from its similarity to
the one described by Goethe, Metam. der Pflanzen.

1 «Bull. Soe. Bot. Fr.,’ tom. iii, 1856, p. 479.
10

146 ALTERATION OF POSITION.

cap. 16, sect. 105; but in that instance median pro-

lification also existed. For my specimens I am in-
debted to Mr. T. Moore.

Fic. 66.—Flower of Dianthus sp., calyx removed; petals turned down
so as to show the stalked flower-buds springing from their axils.

The pistil, too, is necessarily subject to very grave
alterations when affected with this malformation. It
is separated into its constituent carpels; and these
assume a leaf-like aspect, and are in the great majority
of instances destitute of ovules. Indeed, virescence
or chloranthy is very intimately connected with this
aberration, as might have been anticipated, for if the
parts of the flower assume more or less of the condition
of stem-leaves or bracts, it 1s quite natural to expect
that they will partake likewise of the attributes of
leaves, even at the expense of their own peculiar func-
tions.

It occasionally happens that an adventitious bud
arises from the axil of a monocarpellary pistil. This
takes place sometimes in Leguminose, and seems to
have been more frequently met with in T7ifoliwm repens
than in other plants. The species named is, as is well
known, particularly subject to a reversion of the outer
whorls of the flower to leaves, and even to a leaf-like
condition of the pistil. There are on record instances
wherein a leaf-bud has been placed in the axil of a

PROLIFICATION OF THE FLOWER. 147

more or less leaf-like carpel; while at other times a
second imperfect carpel has been met with in the axil
of the first. I have myself seen numerous imperfectly
developed cases of this kind.

It may be asked whether such cases are not more
properly ‘referable to central prolification—whether the
axis is not in such flowers terminated by two, rather
than by one carpel? It is, however, generally admitted
by morphologists that the solitary carpel of Leguminose
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 shgehtly higher up—in fact, in the axil
of the primary carpellary leaf. Griffith figures and
describes* an instance of the kind im a species of
Melilotus. The stalk of the ovary is mentioned as
haying 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
leaf.

' «Linnea,’ vol. xv, p. 266, c. ic. Caspary, ‘Schriften d. Physik.-Oek.
Gesell. zu Konigsberg,’ bd. ui, p. 5, tab. 1, fig. 39, &e.

? Lindley, ‘Veg. King.,’ p. 545; also Clarke on the Position of
Carpels, Linn. Soc.,’ December, 1850. ‘ Proc. Linn. Soc.,’ ii, p. 105.

3 * Notule,’ vol. i, Dicot. p. 127. ‘ Atlas,’ pl. xlii.

OF POSITION.

ALTERATION

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PROLJ FICATION OF THE FLOWER. La)

Complicated prolification—I*rom what has been before
stated it may be seen that prolification of two or more
kinds may coexist in the same flower. Mixed leafy
and floral prolification is not unfrequent in proliferous

‘WW

1 if
inh

g

Fie. 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

b52, ALTERATION ‘OF POSITION.

second flower, or even by a cluster, as is well shown im
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 ehanges were
_ observed :—the swollen portion below the calyx, the
“hip,” was entirely absent; the sepals were leaf-lke
in aspect, the petals unaffected; above the petals
the axis was prolonged for a short distance and then
bore a eirclet of miniature, sessile roses, desti-
tute, indeed, of calyx, but provided with numerous
petals, stamens, and pistils, Above these lateral

XQ MP

jf

Fic. 68.—Rose exhibiting median, axillary, lateral, floral, and leafy
prolification in same flower.

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

PROLIFICATION OF THE FLOWER. 153

by a whorl of five perfect leaves, beyond which, again,
the axis was prolonged into a leafy shoot terminated
by a flower bud, the whole constituting a remarkably
complicated admixture of elements belonging to the
flower, the bud, the inflorescence, and the _leaf-
shoot.’

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 18 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

Fic. 69.—Proliferous Orchis. Diagram showing the arrangement of
the several organs in the seven outer circles of the flower. Each whorl
is numbered, and the position of the axillary buds shown by the small
circles.

1 Moquin-Tandon gives the following references to cases of proliferous
roses, but some I have not been able to verify. ‘Journ. des Sav..,’
22 Mai, 1679. Hottinger, ‘Hphem. 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. Soe. Ital.’ t. v, p. 488. See
also at the end of this section for numerous other references.

154 ALTERATION OF POSITION.

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

x

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t

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

1. The primary flower, with no median bud, the position of which,
had it been present, is shown by the dotted line.

2. Two axillary buds proceeding from 1, and themselves giving
origin to

3, 3. Median buds, and 3’, 3’, axillary buds.

4! 4', Axillary buds, proceeding from 3. No median bud is produced
from 3; its situation, had it been present, is indicated by the dotted
line.
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, AY,

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

On the general subject of Prolification in flowers, in

' «Bull. Acad. Belg.,’ t. xx, part 11, p. 271. See also Bellynck, ‘ Bull.
Soe. Bot. Belg.,’ t. vi, ex. ‘Bull. Soc. Bot. France,’ t. xiv, 1867, Rey.
Bibl., p. 241. Orchis ustulata.

~

PROLIFICATION OF THE EMBRYO. 155

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

Linnzus, ‘ Prolepsis,’ §§ vi et vii. Goethe, ‘ Versuch. Metamorph.,’
eap. xv and xvi, §§ 103—106. Moquin-Tandon, ‘ El. Ter. Veg.,’ p. 362,
&e. Engelmann, ‘ De Antholys.,’ §§ 52—62, &e. Cramer, ‘ Bildungsab-
weichungen, &c. Orchidacew, Umbellifere, Composite, Leguminose,
Primulacee, Rawunculacee. Fleischer, ‘ Missbild. Cultur Gewachs.’

ili, tabs. 3, 4, Rosa. Kirschleger, ‘ Flora,’ 1845, 613, Dianthus, Rosa.
‘ Institut.,’ 1841, No. 413, p. 421, Tragopogon. Baron de Melicoq.,
* Ann. Se. 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.,’ liv. iii, cap.
3, p. 303, pl. xii, f. 306, Rosa. Caspary, ‘ Bull. Soc. Bot. Fr.,’ vol. vi,
1859, p. 235, Rev. Bibl., Pyrus. Eichler, ‘ Flora, 1865, tab. ix, Cleome.
Lindley, ‘Elements of Botany,’ p. 63, &e., Rosa, Epacris, Anagallis,
Pyrus. Irmish, ‘ Flora,’ 1858, p. 38, Pyrus ; and ‘ Bot. Zeit.,’ xix, 1861,
p. 342, Hyacinthus. Duchartre, ‘Bull. Soc. Bot. France,’ 1861, p. 451,
Rosa. Weber, ‘ Verhandl. Nat. Hist. Verein. Rhein. Preuss., &.’ 1858
et 1860. Landrin, ‘Mem. Soc. Sc. Nat. Seine et Oise,’ 1866 2! 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

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

156 ALTERATION OF POSITION.

position. In the latter case, as often happens im the
seeds of the orange, the new products are placed by
the side one of another."

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

CHAPTER, IIT.

HETEROTAXY.

Unpir 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 imtended 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 im
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 moreoyer 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 im 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

1 * Hl. Ter. Veg.,’ p. 364, Adnot.

ADVENTITIOUS ROOTS. 157

considered to come within the scope of teratology, or,
at least, where their production is the result of injury
or of some unfavorable condition to which the plant
is exposed.

Thus the production of adventitious roots on the
stem of the vine is considered to be due to untoward
circumstances impairing the proper action of the ordi-
nary subterranean roots. So, too, the formation of
roots on the upper portions of stems that are more or
less decayed below, as m 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

“ Quin et, caudicibus sectis, mirabile dictu,
Truditur e sicco radiz oleagina ligno.”— Georg.’ Bk. ii.

I have seen many specimens of adventitious roots
produced on the olive in the way just mentioned.

In the ‘Gardeners’ Chronicle,’ January 8th, 1853, p.
21, is described a curious formation of roots in the fissure
between two divisions of alaburnum stem. In the same
journal, January Ist, 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 familar 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

158 ALTERATION OF POSITION.

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 mcreasing weight of branches and
foliage, is also a famihar 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,’

= a = ~

1 ‘Gard. Chron.,’ 1852, p. 51.

ADVENTITIOUS ROOTS. 159

and also in a cabbage where a snail “haying gnawed
a hole into the middle of a leaf at its junction with
the stem, a fascicle of roots was formed, bursting
through the tissue lining the cavity, and covered with
abundant delicate hairs after the fashion of or dinary
radicles.”

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

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

fested on the cotyledons; thus Irmisch describes cases
of this kind in the cotyledons of Buniwm creticwm and
Carum Bulbocastanum.’' 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, 7. 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.

160 ALTERATION OF POSITION.

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

Fries. 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 ovadually i 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 ae
and thus secure a shortened plant.

On the subject of adventitious roots, &c., reference may be made to
Trécul, ‘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 He cidcnae 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.e.,

ADVENTITIOUS BUDS. 16]

Pyrus Japonica, Anemone Japonica, &e. What are
termed suckers, owe their origin to buds formed in this
situation.

If roots be exposed or injured, they will frequently
emit buds. The well-known experiment of Duhamel,
in which a willow was placed with the branches in the
soil and the roots in the air, and emitted new buds from
the latter and new roots from the former, depended on
this production of adventitious organs of either kind.

Gardeners often avail themselves of the power that
the roots have of producing buds to propagate plants
by cuttings of the roots, but m 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." From
the figure it appears as if the lower portion of the root
had been split almost to the extremity, while the upper
portion seems to have a central cavity passing through
it. From the angle, formed by the spht segments
below, proceeds a tuft of leaves, some of which appear
to have traversed the central cavity and to have
emerged from the summit, mingling with the other
leaves in that situation. The production of a flower-
bud has even been noticed on the root of a species of
Impatiens.

Formation of shoots beneath the cotyledons—The tigellar or
axial portion of the embryo plant, as contrasted with
the radicle proper, is very variously developed in
different cases; sometimes it 1s 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, Huphorbia

«Tijdschrift vooor Natuur. Geschied,’ 1836, vol. iii, tab. vii, p. 171.
11

162 ALTERATION OF POSITION.

peplus, and other species, Linaria vulgaris, some Um-
bellifere, &e.'

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 m

Fic. 75.—Leaf proceeding from Fic. 76.—Leaves proceeding
hip of the Rose. from the ovary of Nymphea.

Crategus tanacetifolia, roses, pears, gooseberries, &c.
In a specimen of Nymphea alba T 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 Reeper, ‘ Enum. Euphorb.,’ p. 19. Bernhardi, ‘ Linnea,’ vii, p. 561,

tapes, t..1,.° Wadler, * Subcotyled. sprossbildung, ” © Flora,’ 1850, p.
337. Hooker, ‘Trans. Linn. Soc.,’ vol. xxiv, p. 20 (Welwitschia).

Ee

ADVENTITIOUS LEAVES. 163

For other illustrations of increased leaf-formation,
see Multipheation of foliar organs.

Production of leaves on a usually leafless inflorescence——'l'he
development of the bracts of an inflorescence to such
an extent that 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-

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

164 ALTERATION OF POSITION.

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 Sam-
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 Sempervivum
sediforme,' 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, Taraxracumh, some-
times offers an illustration of this, and also the daisy
(Bellis)? In a specimen of fasciated cowshp 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
1s, 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 versd,
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
shehtest mdication 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.

2 See also Carriére, ‘Revue Horticole,’ 1866, p. 442; and as to pears,
Radlkofer in ‘ Bericht iiber die Thatigkert der Baierischen Gartenbau
Gesellschaft,’ 1862, p. 74, t. 1.

ADVENTITIOUS LEAVES. 165

or any absolute increase in the number of organs, as
in Prolification. Such a case is represented in fig.
78, which shows a portion of the stem of a species of
Valeriana, bearing at the summit, not an inflorescence,
but a tuft of leaves without the slightest indication of
flowers.

Drs. Hooker and Thomson relate that in Northern
India the flowers of Anemone rivularis are very generally
absent, and their place supplied by tufts or umbels of
leaves.’ In the collection of the late Mr. N. B. Ward

Fie. 78.—Tuft of leaves replacing the inflorescence in a species of
Valeriana.

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 Composite,
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. 28.

166 ALTERATION OF POSITION.

Peru, in which the capitula, instead of consisting of
florets, as usual, contained tufts of lmear ciliolated
bracts within the involucre, without a trace of flowers.

In the eleventh volume of the Linnea,’ 1837, p. 301,

Von Cesati figures and describes an analogous case in
Carduus crispus. The same author’ records a similar
instance in the umbel of Seseli coloratwm, 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 Bryophyllum
proliferum. In Digitalis purpurea a similar anomaly
is sometimes met with.

In the apple I have observed leafy shoots bearmg
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 1 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 Phytewma
spicatwm 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 amuch branched panicle, bearing a quantity
of minute bracts, in the axils of which nestle numerous
small buds. In another specimen the inflorescence

' * Linnea,’ xi, p. 301.

ADVENTITIOUS LEAVES—SCALES. 167

preserves its usual catkin-hke shape, but the flowers
are replaced by little tufts of leaves. M. Germain de
Saint Pierre mentions a case wherein the flowers of
Alisma parnassifolia were completely replaced by leaf-
buds."

Here, also, may be mentioned the curious agerega-
tions of scales which occur in some grasses, in Resti-
ace, Juncacee, and other orders, in which the inflor-

Fic. 79.—Spikelets of Fia. 80.—Rose Willow, Salix. sp.

Willedenovia, composed
entirely of scales to the \

exclusion of flowers.

* «Bull. Soc. Bot. Fr.,’ 1856, p. 53.

168 ALTERATION OF POSITION.

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.'

‘** 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 imsect 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-
timuance rather than of the arrest of vegetative develop-
ment.” 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.’

Viviparous plants—'l'he spikelets of certam 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
elumes are changed, sometimes, however, even the
outer and inner palez are wholly unchanged, while
there is no trace of squamule 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, que in olld antea posita, quotannis floruit et
fructus protulit, deinde deponamus in uberiori terra calidi caldarii,
proferet illa per plures annos multos ac frondosos ramos, sine ullo
fructu. Id quod argumento est, folia inde crescere, unde prius enati
sunt flores; quemadmodum vicissim, quod in folia nune succrescit, id,
natura ita moderante, in flores mutatur, si eadem arbor iterum in ollé
seritur.’—Linneus, ‘ Prolepsis,’ § iii.

® * Rev. Hortic,’ May, 1868, ‘Gardeners’ Chronicle,’ 1868, pp. 572, 737.

VIVIPAROUS PLANTS. 169

The grasses most commonly affected in this manner
are Dactylis glomerata!, Poa bulbosa!, Poa annua !,
P. trivialis !, pratensis !, alpina !, angustifolia, and laa,

Fie. $1.—Portion of panicle of Aira vivipara and separate floret.

Cynosurus cristatus, Festuca nemoralis, F. ovina!, Gly-
ceria flutans !, Gl. aquatica, Aira alpina !, ceespitosa!,
Phlewm phalaroides, Lolium perenne!, Alopecwrus pra-
tensis !, Agrostis alba, Holeus mollis !

From an examination of the structure of viviparous
erasses Von Mohl was led to the conclusion that the
lower palea is to be considered as a bract, and not a
perianthial leaf, because the base of the palea sur-
rounds the stem or axis of the spikelet entirely, and
both its margins cohere towards its lower extremity."

A similar condition occurs not infrequently in
Polygonum viviparum, and in Juncacew, Cyperacee, &c.

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

|

170 ALTERATION OF POSITION.

In the genus Alliwm an analogous formation of little
buds or bulbils takes place in heu of flowers; this is
specially the case with A. vineale, the flowers of which
are rarely seen.

Other illustrations of a similar character, where the
adventitious leaf-buds are mixed in amongst the flower-
buds, are cited under the head of Prolification of the
Inflorescence.

Formation of buds on leaves—The formation of litle
bulbs upon the surfaces or edges of leaves, forming
what are called viviparous leaves, has long been familiar
to botanists amongst Alhums. Professor Alexander
Braun,' 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 im
many ferns, in Nymphea guineensis, some Arads, &e.
The same condition has been met with as a teratological
occurrence in the leaves of Cardamine pratensis, Hya-
cinthus Pouzolsii, Drosera intermedia,’ Arabis pumila,
Chelidoniwm majus, Chirita sinensis, Episcia bicolor,*
Zamia, &e.’ 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 hkewise, 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.

2 Naudin, ‘ Ann. Sc. Nat.,’ 2nd ser., 1840, vol. xiv, p. 14, fig. 6, pl. i
(Drosera). St. Hilaire, ‘Comptes Rendus,’ ix, ‘p. 437.

3 Hance, ‘ Hook. Journ. Botany,’ 1849, vol. i, p. 141, pl. v.

4 Booth, ‘ Gard. Chron.,’ Jan. Ist, 1855, P. 4

: Lindley, ‘ Theory of Horticulture,’ ed. 2, p. 273.

6 ‘Hook. Journ. of Botany,’ 1852, iv, p. ’ $06. See also the curious
Begonia gemmipara, ‘ Hook. fil. Tllust. Himal. Plant.,’ t. xiv.

ADVENTITIOUS BUDS. 171

Bryophyllum calyeinum ; Weinmann! figures an instance
of this kind in Alchemilla minima, or they may occur
upon the lower surface of the leaf, as in Ornithogalwm
scilloides and longe-bracteatum. M. Duchartre? 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 Drosera before mentioned.

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

Gardeners occasionally avail themselves of this for-
mation of buds from leaves to propagate plants, e. 4.
Hoya, Gesnera, Gloxinia, &e.

Formation of buds in the pith—This is said to be a

7 1 «Phytanth.,’ n. 36, d.
‘Ann. Science. Nat.,’ 5rd series, 1853, vol. xix, p. 251, tab. 14.

172 ALTERATION OF POSITION.

normal condition in the curious Stangeria paradoxa,'

and Mr. Berkeley records an instance of this in sea-

Fig. 83.—Adventitious buds in sea kale.

kale’ (fig. 83) where the crown had been injured, and
buds were seen sprouting from its centre.
It will be remarked that the adventitious production

Fic. 84.—Hyacinth bulb cut Fie. 85.—Showing the formation
across to induce the formation of of new bulbs on the cut edges of
new bulbs. an old hyacinth bulb.

' Carriére, ‘ Revue Horticole, 1868, p. 184.
? ‘Gard. Chron.,’ 1858, p. 556.

ADVENTITIOUS BUDS. 173

of buds, like that of roots, is very often consequent on
decay or injury. he Dutch bulb-growers have availed
themselves of this latter circumstance in the propa-
gation of hyacinths. Mr. Fortune, who published
some articles on this subject in» the ‘ Gardener’s
Chronicle,’ describes two special modes as adopted by
these skilful horticuiturists—the one to make two or
three deep cuts at the base of the bulb, destroying the
nascent flower-stalk when, after a time, small bulbs are
formed along the edges of the cut surfaces (figs. 84,
85). The other method is effected by scooping out
the interior of the base of the bulb, thus leaving 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. 86).

Fic. 86.—Showing the production of small bulbs on the inner surface
of the scooped-out bulb of hyacinth.

For the formation of supernumerary leaves on the
surface of the normal one, see Multiplication and
Hypertrophy.

Production of gemme in place of spores—An instance of

1863, p. 556, &e.

174 ALTERATION OF POSITION.

this is recorded by Dr. Montagne’ in the case of a
moss, Hncamptodon perichetialis, im which, in the in-
terior of the capsule, in leu of spores numerous minute
gemme 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 ftuseus, in Helwingia, Hrythrochiton
hypophyllanthus, and a few other plants, 1s, of course,
not really to be considered in the light of an actual
growth from the leaf, and it is very doubtful im 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 1s un-
certain at present where it properly belongs.

This curious plant has been described and figured

Fie. 87.—Three-lobed end of Fie. 88.—Three spikelets of
outer palea of Nepaul barley Nepaul barley.
bearing supplementary florets.

1 * Ann. Nat. Hist.,’ 1845, vol. xvi, p. 355.

175

ADVENTITIOUS FLOWERS. (4

by Irmisch in the 13th volume of the ‘ Linnea,’ p. 124,
t. iv; also by Professor Henslow, ‘ Hooker’s Journal
of Botany,’ 1849, vol. i, p. 33, tabs. 2, 3. The lower
palea of this plant forms an inverted flower-bud upon
its midrib. In some fresh specimens which I have

Fic. 89.—Lip of outer palea Fie. 90.—Supplementary rachil-
of Nepaul barley. lus or outer palea of Nepaul bar-
; ley bearing florets.

Fie. 91.—Diagram showing ar- Fic, 92.—Supplementary floret
rangement of supplementary ra- of Nepaul barley; palea removed.
chillus and florets.

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

176 ALTERATION OF POSITION.

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 palez in
the primary flower are sometimes extended into a long
awn. <A similar awn may also be occasionally found
on the tips of the pales 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.’

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,*
and this is effected in various ways, as by continuous
**pinchine”’ or rer.oval of the leaf-buds, by pruning,
ringing the bark, confining the roots, limiting the

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

? 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.

HETEROTAXY. i

supply of nutriment, and other means all based on the
same principle. Some of the Cape bulbs (Cyrtanthus)
are known not to produce their flowers till their leaves
have received, In some manner, a check. Fires which
often destroy the herbage thus have the effect of 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—'hat the spine, as a con-
tracted branch, should occasionally produce fiowers is
not to be wondered at, though the occurrence is by no
means common. M. Baillon showed at a meeting of
the Botanical Society of France (‘ Bulletin,’ vol. v, 1858,
p- 316) a branched spine of Gleditschia bearing a flower
at the end of each of the 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
‘Gardeners’ Chronicle’ for 1865, p. 769, by the Rev.
M. J. Berkeley, who describes the formation of
flower-bud on the surface of a petal of Clarkia elegans.
Reasoning from analogy there seems no reason why
buds should not be formed on the petals as well as on
the leaves.

12

178 ALTERATION OF POSITION.

Formation of buds on fruits—'l'his is a pomt of some
moment with reference to the share which the axis
takes im the production of ‘‘inferior” fruits. A very
frequent malformation in pears is one wherein a
second pear proceeds from the centre of the first, and
even a third from the centre of the second.’ 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.” Further
reference to these fruits will be made under the head
of Hypertrophy.

M. Trécul 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.’ A similar growth
was observed in the fruit of Philadelphus speciosus by
M. A. Gris, who observed that the so-called calyx-tube
was provided with two small bracts, from the axil of
one of which proceeded a small flower-bud.*

The fruits of Opuntia Salmiana, O. fragilis,’ O. mona-
cantha, and of some species of Hchinocactus, have been
observed to form small fruit-like branches around their
summits. M. Napoléon Doumet describes the fruit as
ripening as usual, but as being destitute of seeds m 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;
Irmish, ‘ Flora,’ 1858, p. 38, &c.

* Caspary, ‘ Bull. Soc. Bot. Fr.,’ vol. vi, 1859, p. 235; also Payer, ibid.,
vol. i, 1854, p. 283.

3 Trécul, ‘ Ann. Sc. Nat.,’ 2nd ser., vol. xx, p. 339.

4 «Bull. Soc. Bot. Fr.,’ vol. vii, 1858, p. 331.

* «Bull. Soe. Bot. Fr.,’ vol. 1, p. 306, vol. v, p. 115. ‘ Illustr. Hortie.,’
xii, 1865, Mise. 79. ‘ Rev. Horticole,’ 1860, p. 204, et 1867, p. 43.

HETEROTAXY. 179

and then a circle of small buds, like those of the stem,
may be seen at the top of the fruit, each bud springing
from the axil of a little tuft of wool and spines found
on the fruit. These little buds elongate into long
shoots, produce flowers the following year, which
flowers exhibit the same peculiarity. Gasparini and
Tenore are said to have recorded the same fact as long
since as 1832. The specimen from which the figure (fig.
93) was taken produced its fruits inthe Royal Gardens
at Kew, and is now preserved in the museum of that
establishment. ‘The adventitious growth in these cases

Fie. 95.—Small buds projecting from the edges of the fruit in Opuntia.

appears to arise from the tufts of spies, which, it has
been suggested, are the homologues of the sepals.
There can, however, be little doubt that the outer and
lower portion of the fruit of Opuntia and its allies is a
dilatation of the flower-stalk. This is borne out by
the fruits of Pereskia, which bear leaves on their sur-
face arranged spirally ; indeed, the fruits of Pereskia
Bleo are mentioned as producing buds from their

180 ALTERATION OF POSITION.

summits, in the same way as the Opuntia just cited.
P. Bleo is said, by M. Delavaud,' 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 sprmgs
from the upper portion of the peduncle which is dilated
to invest the true carpels. In other instances it 1s due
to an adhesion of the pedicel to the side of the fruit. In |
either case the production of an adventitious bud might
be considered as an illustration of prolification of the’
inflorescence, though not as was supposed by Moquin
and others of axillary prolification.’

Buds have also been produced artificially on the
surface of some of the fruits in the construction of
which the axis is supposed to share; thus, the unripe
fruits of some species of Lecythis were stated by Von
Martius, at a meeting of the German Naturalists at
Carlsruhe, to produce buds when placed in the earth.
The fruit of these plants is probably of the same nature as
that of the Pomacew, and Baillon® succeeded in producing
buds on the surface of the inferior ovary of Jussica.

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
question.

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

' «Bull. Soe. Bot. Fr., 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. Fr.,’ vol. v, 1855,
p. 672. Seringe et Heyland,‘ Bull. Bot., i, p. 8. ‘Pallas Enum, Plant.
Hort. Demidoff,’ append. e, ic.

3 « Adansonia,’ i, 181.

HBETEROTAXY. 18]

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-
cation strictly.

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

Fiq. 94.—Distended pod of Sinapis arvensis bearing in the interior
stalked fiower buds.

two flower-buds were found projecting from a raised
central line, cérresponding, as it. would seem, to the
midrib, and not to the margins of the carpel. Similar
cases have occurred in Nasturtiwm amphibium, Brassica
Rapa, and Passiflora quadrangularis.

In Bromfield’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 bemg opened, were
found to contain petals of the usual colour, which in
the pods above had burst from their confinement and
appeared as semi-double fiowers ; the valves of the pod
answering to the true calyx. * * * From their verti-
cillate arrangement it is evident that these petaloid
expansions were not transformed seeds, but simply a
development of the common axis within the ovary into
an abortive whorl of floral organs, besides which there
were evident rudiments both of stamens and germens

182 ALTERATION OF POSITION.

in the centre of the bundle.” Baillon' also records a
case of the same nature in Simapis 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

“Sin

" r vie

Fia. 94.— Portion of the interior Fia. 95.—Adventitious pod from
of thesilicle in Cheiranthus Cheiri, fig. 94, enlarged.
showing adventitious pod in the
place of an ovule.

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 Beckea diosme-

! * Adansonia,’ vol. iii, p. 351, tab. xii.

ne

HBTRROTAXY. 183

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

Fie. 96.—Section of Barbarossa Fie. 97.—Grape with supple-
grape showing adventitious grape mentary fruit in the interior.
in the position of a seed.

In the normal flower there is a turbinate hollow
calyx, whose limb is divided into five serrated lobes ;
alternating with these latter, and springing from the
throat of the calyx, are the petals. Originating from
the same annular disk as the petals are the stamens,
seven or eight mm 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

184 ALTERATION OF POSITION.

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 carpellary
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 Bechkea there were perfect stamens proceeding

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

HETEROTAXY, 185

from perfect and completely closed ovaries. Moquin-
Tandon’ cites from Agardh an instance which seems
more closely to resemble the state of things in the
Beckea, and which occurred in a double hyacinth,
wherein both anthers and ovules were borne on the
same placenta. Probably, though the fact is not
stated, the ovary of the hyacinth was open; and we
are told that the flower was double—that it was, in
fact, modified and changed in more organs than one;
while in the Beckea 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 alhed Babing-
tonia, a prolongation of the placenta.’

Formation of pollen within the ovules—This has now been
recorded in two instances by Mr. 8. J. A. Salter in
Passiflora cerulea and in P. palmata,’ and by the author
in Rosa arvensis.*

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

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

' «Hilém. Térat. Végét.;’ p. 218.

* 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.

186 ALTERATION OF POSITION.

less split open at the distal end, indicating a tendency
towards dialysis. The pollen-bearmg 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-lke
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

POLLINIFEROUS OVULBES. 187

of the ovule, while the pollen was formed within the
nucleus. In no case was any trace of embryo sac
to be seen.

The main interest, as Mr. Salter remarks, in these
cases is physiological; so far as structure alone is con-
cerned, there does not appear any reason why pollen-
erains 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-

188 ALTERATION OF POSITION.

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 imstances 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. Hven 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 im any
future alleged occurrence of the latter phenomenon to
ascertain whether any or all of the apparent ovules are,
or are not, anthers in disguise.

Homomorphic flowers of ‘* Composite.’ '—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. m the Chrysanthemum, Dahlia, &c. &e., the
florets are all hgulate. 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 Liguliflore, 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

HETEROTAXY. 189

deviations from the customary arrangement of the in-
florescence, such as the change from a definite centri-
fugal form of inflorescence to an indefinite centripeta!
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 twice 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
anticipée,”’ 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
er branches usually sterile become fertile ; this happens
in Hquisetacee,' in Restiacew, and other orders. In the
equisetums, the condition in question has been specially
noticed to occur after prolonged drought.

Hawisetacee 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 Saliv cinerea were placed at the ends of the

! Duval Jouve, ‘ Hist. Equiset. France,’ 1864, p. 154.

190 HETEROGAMY.

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

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 marginal 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: Scolo-
pendrium vulgare, Polypodium anomalum, Hook., As-
plenium Trichomanes, Cionidium Moorei.’

CHAPTER IV.
HETEROGAMY.

Tis 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.

> Moore, ‘ Nature-Printed British Ferns,’ 8vo edition, vol. ii, p- 135,
tab. Ixxxv, B, Ke.

HETEROGAMY. 19]

tually suppressed, a tendency towards structural com-
pleteness rather than the reverse. It must be also
understood that 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
that plants hermaphrodite as to structure are by no
means necessarily so as to function.

The simplest case of this alteration in the relative
position of the sexes is that which occurs in moncecious
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 moneecious plant. Cultivated
maize, Zea Mays, frequently exhibits alterations of this
kind; under ordinary circumstances, the male inflor-
escence 1s 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.’ 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 Conifere 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

' 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.

192 HETEROGAMY.

in July, 1860, some malformed cones of Abies 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-lke crest had become more
elongated and pointed, in fact,
more or less resembling the or-
dinary bracts." Mohl, Schleiden,
and A. Braun have observed
similar cones in Pinus alba, and
Cramer figures and _ describes
androgynous cones in Lariv
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.

Fie. 100. — Spike of
Carex acuta, with female On the subject of this section the reader
flowers at the summit. may consult A. Braun,. ‘ Das Individ.,’ 1853,
p. 65. Caspary, ‘De Abietin. flor. fem.
struct. morphol.’ Schleiden, ‘ Principles,’ English edition, p. 299. Mohl,
‘Verm. Schrift.,” p. 45. Meyen in ‘Wiegm. Archiv.,’ 1838, p. 155.
Cramer, ‘ Bildungsabweich,’ p. 4, tab. v, figs. 18—17. Parlatore, ‘ Ann.
Se. Nat.,’ ser. iv, vol. xvi, p. 215, tab. 13a. See also under the head of
Prolification, Substitutions, &e.

1 Professor Dickson concludes from the examination of these strue-
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.

HETEROGAMY. 193

Change from the monecious to the diwcious condition—'lhis
is of less frequent occurence than might have been
anticipated. In the ‘Gardeners’ Chronicle,’ 1847,
pp. 541 and 558, several instances are noted of walnut
trees bearing female flowers to the exclusion of males.
The mulberry tree has also been noticed to produce
female blossoms only, while in other plants male
flowers only are developed.

It seems probable that the age of the plant may
have something to do with this production of flowers
of one sex to the exclusion of the other.

Change from the diccious to the monecious condition.—Andro-
gynism—T'his is of far more common occurrence than
the preceding.

Fie. 101—Monecious inflorescence of Hop.

In the hop (Humulus LInpulus), when moncecious,
the female catkins are usually borne on the ends of

the branches as shown in the cut (fig. 101), and a similar
| 13

194. ALTERATION OF POSITION.

thing has been noticed in Urtica dioica by Clos, ‘ Bull.
Soc. Bot. France,’ vol. 9, p. 7.

Baillon (‘ Etudes du groupe des Euphorbiacées,’ p.
205) mentions the following species of that order as
haying been seen by him with moncecious inflorescence:
Schismatopera distichophylla, Mozinna peltata, Hermesia
castaneifolia. Oliver mentions (‘ Hook. Icon. Plant..,’
t. 1044) that in Leitneria floridana the upper scales of
the male catkin occasionally subtend an ovary.

It would seem that external conditions have some
effect in determining the formation of one sex, as in
some species of Carex, while in the case of Salix repens,
Hampe' 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’ says that a plant of Stauntonia latifolia
which for some years produced stamens only, now
produces flowers of both sexes; it was dicecious, but
is now moncecious. 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.* In addition to the genera, already
named, in which this production of flowers of both
sexes has been observed may be mentioned Tawus!
Guinera! Urtica! Mercurialis! Restio! Cannabis !
Saliz! Humulus! as well as others in which the
change is less frequent.

Among cryptogams a similar change occurs. As an
illustration may be cited Leucobrywn gigantewm, as
quoted from Miillerin 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 «Linnea,’ xiv, 367.
? ‘Rev. Hortic.,’ January, 1867.
5 See Royle, ‘Man. Materia Medica,’ ed. 1, p. 567.

UNISEXUALITY. L95

Berol.,’ ii, p. 259. Tausch, ‘ Bot. Zeit.,’ 1833, i, p. 229. Koch, ‘ Synops.
Flor. Germ.,’ 740. Host, ‘Flor. Aust.,’ ii, p. 641 (8. mirabilis). See
also Hegelmaier, ‘Wiirttemberg Naturwissenschaft Jahreshefte,’ 1866,
p. 30. Other references to less accessible works are given in ‘ Linnea,’
xiv, p. 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 since the
demonstration of the fact by Darwin and others, that
many plants, structurally hermaphrodite, require for the
full and perfect performance of their functions the co-
operation of the stamens and pistils, belonging to dif-
ferent individuals of the same species.

Some of the Ranunculacew constantly exhibit a ten-
dency towards the dicecious condition, and the rarity
with which perfect seeds of Ranunculus Ficaria are
formed is to be attributed, in great measure, to the
deficiency of pollen in the anthers of these flowers.
Ranunculus auricomus also is frequently sterile. Speci-
mens of Ranunculus bulbosus may be met with in which
every flower is furnished with carpels, most of which
have evidently been fertilised, although there are no
perfect stamens in the flowers.

Knight and other vegetable physiologists have been
of opinion that a high temperature favours the pro-
duction of stamens, while a lower degree of heat is
considered more favorable to the production of pistils,
and in this way the occurrence of ‘‘blind”’ strawberries
has been accounted for. Mr. R. Thompson, writing
on this subject, speaks of a plantation of Hautbois
strawberries which in one season were wholly sterile,
and accounts for the circumstance as follows:
the plants were taken from the bearing beds the
year previous, and were planted in a rich well-manured
border, in which they started rapidly into too great
luxuriance, the growth being to leaves rather than to
fruit. The following season these same plants bore

196 ALTERATION OF POSITION.

a most abundant crop, hence these plants were acci-
dentally prevented from perfecting their female organs.’

Mr. Darwin’ cites from various sources the following
details relating to strawberries which it may be useful
to insert in this place, as throwing some heht 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 plents
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 lmits
of its geographical distribution, and its stamens, on
the other hand, are stated to be more frequently de-
veloped in more southern districts.

Honckenya peploides affords another illustration of the
sexual arrangements in the flower being altered as it
would seem by climatal conditions. Thus, in the
United States, according to Professor Asa Gray, the
flowers are frequently hermaphrodite, while in this
country they are usually sub-dicecious.®

Treviranus* says that the flowers of Hippuris and
Callitriche are apt to be hermaphrodite in summer, but
female only at a later period.

For further remarks on this subject, see sections
relating to suppression of stamens and pistils.

1 Thomson, ‘Gardener’s Assistant,’ p. 577.

2 * Variation of Animals and Plants,’ i, 353.

> Babington, ‘ Ann. Nat. Hist.,’ vol. ix, 1852, p. 156.
“ «Phys. der Gewiichse,’ i, p. 325.

HERMAPHRODITISM. 197

Change from unisexuality to hermaphroditism.—'l'his occur-
rence depends on one of two causes, either organs are
developed (stamens or pistils as the case may be),
wihch are habitually absent in the particular flower ;
or some of the stamens may be more or less com-
pletely converted into or replaced by pistils, or vice versa.

The first condition is the opposite of suppression ;
it is, as it were, a restoration of symmetry, and might
be included under the head of regular peloria, inas-
much as certain organs which habitually undergo
suppression at a certain stage in their development,
by exception, go on growing, and produce a perfect,
instead of an imper rfect 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, diclnous flowers may,
under certain conditions, become perfect, at least
structurally. I have myself seen hermaphrodite
flowers in Cucurbita,’ Mercurialis, Cannabis, Zea Mays,
and Aucuba japonica, as well as in many festiacee,
notably Cannamois virgata and Lepyrodia hermaphro-
dita. Spinacia oleracea, Rhodiola rosea, Cachrys taurica,
and Hmpetrum nigrum are also occasionally herma-
phrodite.

Gubler* alludes to a similar occurrence in Pistacia
Lentiscus, wherein, however, he adds that there was a
deficiency of pollen i in the flowers.

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

Baillon* enumerates the following Huphorbiacee as
having exceptionally produced hermaphrodite flowers,

1 See also Schlechtendal, ‘ Linnea,’ viti, p. 623, and Lindley, ‘ Veg.
Kingd.,’ p. 315.
“Bull. Soc. Bot. France,’ vol. ix. p. 81.
3 Cited in Henfrey, ‘Bot. Gazette,’ 3, p. 11.
4 Baillon, ‘Etudes du Groupe-des Euphorbiacées,’ p. 205, tab. xv,
fig. 19, tab xix, fig. 31.

198 ALTERATION OF POSITION.

Crozophora tinctoria, Suregada sp., Phyllanthus longi-
folius, Breynia sp., Philyra brasiliensis, Ricinus com-
munis, Conceveiba macrophylla, Cluytia semperflorens,
Wall. non Roxb. Mercurialis annua and Cleistanthus
polystachyus.

In some of these cases the hermaphroditism is due
to the development of anthers on the usually barren
staminodes, though, in other cases, the stamens would
seem to be separate, independent formations, as they
do not occupy the same relative position that the
ordinary stamens would do if developed."

Robert Brown’ observed stamens within the utricle
of Carex acuta, and Gay is stated by Moquin (‘ El. Ter.

Fie. 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, ‘Mém. Soc. Nat. Hist. Paris,’ I, p. 16; herma-
phrodite flowers in Euphorbia esula.
2 * Prod. Flor. N. Holl.,’ p. 242.

HERMAPHRODITISM. 199

Veg.,’ p. 343) to have observed a similar occurrence
in Carex glauca.

Paasch' observed a similar occurrence in C. cespitosa,
and Schauer, in C. paludosa,” though in the latter in-
stance the case seems to have been one of transfor-

“mation or substitution rather than one of hermaphro-
ditism.

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.®

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

Fie. 103—Hermaphrodite flower of Carica Papaya.

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

1 © Bot. Zeit.,’ 1837, p. 335.

2 « Pflanz, Terat.,’ von Moquin-Tandon, p. 208.

3 Schnizlein, loc. cit. 5)

4 «Bot. Mag.,’ tab. 5160, fig. 4. See also ‘Gard. Chron.,’ 1860, pp.
146, 170; 1861, p. 1092. .

200 ALTERATION OF POSITION.

formed, as happened also in a garden variety of a
Fuchsia, wherein, however, the change was less perfect
than in the Begonia, and in which, as the flower is
naturally hermaphrodite, the alteration is of the less
importance.

In hermaphrodite flowers of Carica Papaya (fig. 103)
there is a single row of five stamens instead of two rows
of five each as in the normal male flowers, the position of

Fra. 104.—Ovuliferous anthers—Cuewrbita.

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

CHANGE OF DIRECTION. 201

arrangement of parts in the normal male flowers to
be as follows :

P P P P P

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

18) S

5 P
5 st

5 c

One of the most curious cases of this kind recorded
is one mentioned by Mr. Berkeley,’ 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, ike 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.

CHAPTER V.
ALTERATIONS IN THE DIRECTION OF ORGANS.

Tue 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. Chron.,’ 1851, p. 499.

202 ALTERATION OF POSITION.

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
materially.

Change in the direction of axile organs, roots, stems, &c,—
The roots frequently exhibit good illustrations of the
effect of the causes above mentioned in altering the
natural direction. ‘The roots are put out of their
course by meeting with any obstacle in their way.
Almost the only exception to the rule in accordance
with which roots descend under natural circumstances,
is that furnished by Trapa natans, the roots of which
in germination are directed upwards towards the sur-
face of the water. So in Sechiwm edule, the seed
of which germinates while still in the fruit, the roots
are necessarily, owing to the inverted position of the
embryo, directed upwards in the first stance.

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 Itahan 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, chesnuts, and other
plants which may be met with in the nurseries.

DIRECTION. 903

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.'

Climate appears sometimes to have some influence

' The following details as to the method pursued by Mr. McNab, of
the Edinburgh Botanic Garden, may not be uninteresting in this place.
They are from the pen of Mr. Anderson, and originally appeared in the
‘Gardeners’ Chronicle.’

“The mode of inducing leaders to proceed from laterals is a matter
of comparatively little concern among the generality of deciduous trees,
for they are often provided with subsidiary branches around the leader,
at an angle of elevation scarcely less perpendicular, but the laterals of
all Conifers stand, as nearly as possible, at right angles. Imagine the
consternation of most people when the leader of, say, Picea nobilis, P.
Nordmanniana, or P. Lowii is destroyed.”

In a specimen of the latter plant the leader had been mischievously
destroyed, to remedy which Mr. Mec Nab 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. Lowii
above referred to. Resort must, therefore, be had to one of the best
favoured laterals, but how is it to be coaxed from the horizontal position
of a lateral to the perpendicular position of a leader? The uninitiated
in these matters, and, in fact, practical gardeners generally, would at
once reply, by supporting to a stake with the all-powerful Cuba or bast-
matting. Butno. 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
asupremacy. Singular provision in nature this, which, thanks to the
undivided attention of a careful observer, has been fully appreciated and
utilized.”

f

204 ALTERATION OF POSITION.

on the direction of branches, thus Dr. Falconer, as
quoted by Darwin,’ 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 Lobinia 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 multiphed 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.”

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. 277,
* Quoted in ‘ Gard. Chron.,’ 1867, p. 654.

CHANGE OF DIRECTION. 205

then elongate into a shoot and no cavity is formed. <A
fig observed by Zuccarini (figs. 105, 106) appears to
have been formed in a similar manner, the flower-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—'T'he 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 1s bent downwards the leaves may be placed in the
opposite direction; thus in some specimens of Galiwin
Aparine growing on the side of a cliff from which there
had been a fall of chalk, the stems, owing apparently to

Fie. 105.—Fig showing pro- Fie. 106.—Section of the same.
longed inflorescence and projecting
flowers.

the landslip, were pendent, but the leaves were abruptly
bent upwards.
One of the most singular instances of an inverted

206 ALTERATION OF POSITION.

direction of the leaves is that presented by a turnip (fig.
107) presented to the Museum of King’s College, London,
by the late Professor Edward Forbes. The turnip is
hollow in the interior and the majority of the leaves
springing from its apex instead of ascending into the
hight 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

\
YY \

4
4

\

A WN

\\

Fia. 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-

DIRECTION. 207

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 Gloxinia, the normal
flowers of which are irregular and pendent, there is
now in common cultiy ation a peloriate race in which
the flowers are regular in form and erect in position.

Fie. 108.—Flower of normal Gloxinia.

Fig. 109.—Flower of Glowinia, erect and regular (regular Peloria).

Fig. 108 shows the usual irregular form of Glovinia,
with which may be contrasted figs. 109, 110 and 111.

208 ALTERATION OF POSITION.

Fig. 109 shows the regular erect form; fig. 110
the calyx of the same flower; while in fig. 111 are
shown the stamens and style of the two plants 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.

Fia. 110.—Calyx of erect Fie. 111.—Stamens of erect
Gloxinia. 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-

CHANGE OF DIRECTION, 209

like torus are bent downwards towards the stunted
carpels instead of being, as they usually are, straight.

Amongst orchids, where the pedicel of the flower or
the ovary is normally twisted, so that the labellum 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

Fre. 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' mentions a flower of Rosa alpina in which
two of the petals were erect, while the remaining ones
were much larger and expanded horizontally. The
same author quotes from M. Desmoulins the case of a
species of Orobanche, in which a disjunction of the
petals constituting the upper lip took place, thus libe-

1 Loe. cit., p. 315.
14

210 ALTERATION OF POSITION.

rating the style and allowing it to assume a vertical
direction.

M. Carriere! has described an instance wherein two
apples were joied together, a larger and a smaller one ;
the former was directed away from the centre of the

il
1

= =3 g —-_ en Bi

Fre. 118.—Flower of Cuphea miniata enlarged, showing protrusion
and hypertrophy of an erect placenta, after Morren.

Fia. 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

1 § Rev. Hortic.,’ 1868, p. 110.

CHANGE OF DIRECTION. 211

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 Cissus.’

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

! Planchon and Marés, ‘ Ann. Se. Nat.,’ 5 ser., tom. vi, 1866, p. 228,
tab. xii.

9

2 * Bull. Acad. Belg.,’ xviii, part 11, p. 293.

BOOK II.

DEVIATIONS FROM THE ORDINARY FORM OF ORGANS.

Ivy 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.’ 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 they

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 symmetrical halves (bi-lateral symmetry).
Others, again, have applied the term symmetry to the number of the
parts of the flower, reserving the terms “regularity” or “irregularity”
_ for the form. It is here used in a general sense to express the plan of the

flower, and thus includes the arrangement, form, and number of its
component elements.

214 FORM OF ORGANS.

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 vwice versd. 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 1s 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 followmg

1 See Baillon, ‘ Adansonia,’ v, 176.

FORM OF ORGANS. 215

terms—Stasimorphy, Pleiomorphy, Metamorphy, and
Heteromorphy—to include teratological changes really
or apparently due to one or other of the causes above
mentioned. ‘The classification here adopted is of course
to a considerable extent an arbitrary one and subject
to correction or modification, as the knowledge of the
development of the flowers in the various genera of
plants advances.

PATE ik.

SLAs da OUR Paty

Devtations 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 Yraowc-pwoogware.

STASIS. 217

CHAPTER I.
PERSISTENCE OF JUVENILE FORMS.

Tue retention in adult hfe of a form characteristic
of an early stage of development, and therefore usually
transient, may be manifested in any of the organs of
he 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

Fie. 115.—Juniperus sinensis. Two forms of leaves on branches of
the same shrub.

218 PERSISTENCE OF JUVENILE FORMS.

occasionally happens that the plant continues to form
throughout its existence leaves such as are usuall
produced only i ina young state; thus M. Gubler (Bull.
Soc. Bot., Fr.,’ vol. vin, 1861, p. 527) describes a plant of
Pinus pines i 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 erows 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.’ 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 Stesomy” in
‘Bull. Acad. Belg.,’ t. xix, part u, p. 519.

REGULAR PELORIA. 219

CHAPTER II.
REGULAR PELORIA.

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 Linnzeus, 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.' 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, Fia. 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

' «On the existence of two forms of Peloria,” by M. T. Masters.
‘Nat. Hist. Review,’ April, 1865.

220 REGULAR PELORIA.

in other species of this genus. Baillon,' 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

A
\

“a a

Fic. 118. — Regular _ peloria, Fie. 119.—Double Violet, flower
Viola. regular, petals multiplied, stamens

and pistils petaloid.

| Baillon, ‘ Adansonia,’ iv, p. 149.

REGULAR PELORIA. 221

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-hke organs.’ 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
(Alsodeie) 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 alhed genus Geraniwm; 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. LHven 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. Vern.,’ fol. 4,
f.1,2. Viola martia multiplici flore.

222 REGULAR PELORIA.

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 Tropawolum
majus in Which the flowers were perfectly regular and
devoid of spurs,’ 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 Papilionacee 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 Gesneracee, Bignowacee, Scrophu-
lariacee, and other families of like structure, regular
peloria is not uncommon. se 120 represents a case of

Fic. 120.—Regular peloria, Hecremocarpus scaber.

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

1 ‘Linnea,’ 1837, p. 128.
2 M. Bureau, ‘ Bull. Soc. Bot. Fr.,’ ix, p. 91, describes two genera of
Bignoniacee in which the flowers are normally regular and six parted.

REGULAR PELORIA. 993

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

A solitary flower of Pedicularis sylvatica was found
by the Marquis of Stafford near Dunrobin Castle in
Sutherlandshire, in which the usual ringent form
of the corolla was replaced by the form called salver-
shaped. There were six stamens, four long and two
short. Sir W. Hooker and Mr. Borrer are stated to
have found a similar flower in the same locality in
1809."

The passage of hgulate to tubular corollas among
Composite 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.

Bae. 121.—Flower of Cattleya marginata. Lip replaced by a flat
petal.

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

! See ‘ Trans. Linn. Soc.,’ vol. x, p. 227.

224 REGULAR PELORIA.

character, asin Dendrobium normale, Oncidiwm heteran-
thum, Thelymitra, etc. Fig. 121, reduced from a cut in
the ‘ Gardeners’ Chronicle,’ 1854, p. 804, represents an
instance of this kind in Cattleya marginata.

From the same journal the following account of
a case of peloria in Phalenopsis 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 Selenipediwm
caudatuwm with a flat lip.

M. Gris’ has placed on record some interesting cases
of peloria of this kind in Zingiber zerwmbet ; mm the more
complete forms the andrcecium or staminal series was
composed of six distinct pieces, the three mner of
which were fertile, while in the ordinary flower the
androecium is composed of two pieces, ‘fa lip” and a
fertile stamen. ‘Is it not a matter of regret,” says
M. Gris, “to be obliged to call the latter the normal
flower ?”’

Under this head may likewise be mentioned those
cases in which the normal, or at least the typical 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 Bocconia with a

1 «Ann. Sc. Nat.,’ ser. 4, 1859, tom. xi, p. 264, tab. 3.
2°, Ter. Veg.,’ p. 342.

REGULAR PELORIA. 225

corolla. Arwm maculatum has likewise been met with
provided with a genuine perianth as in Acorus and
other Orontiads. The unusual development of the
sexual organs in diclinous flowers has been alluded to
under the head of heterogamy, and other cases where
the symmetry of the flower is rendered regular, by the
development of parts ordinarily suppressed, will be
found in the chapters relating to deviations from the
usual number of organs.

This change, or rather this persistence of a form that
is usually transient, is generally accompanied by some
other alterations. Change of direction, as has been
already mentioned, is one of the most common of
these ; separation of the petals (Antirrhinum, Verbascum,
&c.), and even their appearance in leaf-like guise, are not
infrequent (Delphinium, Antirrhinum, Verbascum, &c.)
At other times multiplication or increased number of the
whorls of petals takes place, often, but not always, at the
expense of the sexual organs of the flower. Perhaps
even more frequent is the increased number of parts
in the same whorl in cases of regular peloria ; thus, in the
Pelargoniums before alluded to, the parts of the flower
are frequently regulated by the number six instead
of five.

This form of peloria is most generally met with in
flowers that are placed at the end or in the centre of
the inflorescence, or in such flowers as occur singly at
the end of the flower-stalk, as in Tropewolum, Viola, &e.
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-

| 15

226 REGULAR PELORIA.

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

The occurrence both of regular and irregular peloria
on the same plant has frequently been observed in
linaria. It hasalso 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.”

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, Xe.

*Delphinium peregrinum ! Streptocarpus Rexi.
*Nigella damascena ! *Digitalis purpurea.
*Aquilegia vulgaris! *Scrophularia aquatica.
*Viola odorata! *Pentstemon.
hirta. *Linaria vulgaris!
Kpimedium, sp. * Antirrhinum majus !
*Pelargonium zonale! Verbascum nigrum !
3 inquinans! Columnea Schiedeana.
Tropeolum majus! Halenia heterantha.
*Wistaria sinensis. Galeodolon luteum.
Lupinus. Prunella vulgaris!
*Cytisus Laburnum ! Salvia, sp. !
Trifolium repens ! Teucrium campanulatum.
*Composite, gen. pl. ! Betonica alopecuros.
Lonicera, Periclymenum ! Eecremocarpus scaber.

' Marchand, ‘ Adansonia,’ vol. iv, p. 127.
2 * Bull, Acad. Belg.,’ xvii, p. 17. ‘“ Fuchsia,” p. 169.

REGULAR PELORIA. 227

Pedicularis sylvatica, Atriplex, sp.
Zingiber Zerumbet. Cattleya Mossia !
Phalewnopsis amabilis ! marginata.
Phalewnopsis Schilleriana. Calanthe vestita!
Habenaria. Oncidium, sp. !
*Orchis morio. Selenipedium caudatum.
mascula. Arum maculatum.

*Dendrobium, 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
Peloria. —

Giraud, ‘Bot. Soc. Edinb., Dec. 12, 1839, Antirrhinum. Dareste,
‘Ann. Se. Nat.,’? ser. 2, 1842, xviii, p. 220, Delphinium. C. Morren,
‘Fuchsia,’ p. 90, Calceolaria, ‘ Bull. Acad. Belg.,’ xx, part i, p. 57; and
E. Morren, ‘ Bull. Acad. Belg.,’ 2nd ser., xix, p. 224, Glowinia. Richard,
‘Mém. Soe. d’hist. nat.,’ ii, p. 212, tab. 3. Lindley, ‘Journ. Linn. Soc.,’
iii, p. 9, Dendrobiwm. Michalet, ‘Bull. Soc. Bot. France,’ vii, p. 625,
Betonica. Gubler, ‘Bull. Soe. Bot. Fr.,’ ix, 81, ‘Des anomalies aber-
rantes et regularisantes.’ Reichenbach fil. ‘De pollinis orchid. genesi
ac structura,’ 1852, Oncidiwm. Clos, ‘Mém. Acad. Toulouse,’ vi, 1862,
Salvia. Caspary, ‘ Verhandl. Phys. @kon. Gesell. Koénigsberg,’ 1860,
i, 59, Colwmnea. Bureau, ‘Bull. Soc. Bot. Fr.,’ 1861, vol. viii, p. 710,
Streptocarpus. Darwin, ‘ Variation of Animals and Plants,’ u, 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 regularité 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. ;

iPrAs bis

PLEIOMORPHY!

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.

CHAPTER IL.

IRREGULAR PHLORIA.

Tur term peloria was originally given by Linné
to a malformation of Linaria vulgaris, with five spurs
and five stamens, which was first found in 1742 near
Upsal. This was considered so marvellous a circum-
stance that the term peloria, from the Greek védwp, a

1 TI\etog-poppwore.

IRREGULAR PELORIA. 229

prodigy, was applied to it.'| After atime other irregular
flowers were found in-like condition, and so the term
peloria became applied to all cases wherein, on a plant
habitually producing irregular flowers, regular ones
were formed. The fact that this regularity might arise
from two totally different causes was overlooked, or at
least not fully recognised, even by Moquin-T'andon
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
recular or congenital peloria (see chapter on that
subject) ; but where a flower becomes regular by the
increase in number of its irregular portions, as in the
Tinaria already alluded to, where not only one petal
is spurred, but all five of them are furnished with such
appendages, and which are the result of an irregular
development of those organs, the peloria is evidently
not congenital, but occurs at a more or less advanced
stage of development. ‘To this latter form of peloria
it is proposed to give the distinctive epithet of
irregular.

Peloria is either complete or incomplete; it 1s 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 * Ameen. Acad.,’ i, p. 55, t. iii (1744) :—The following note refers to
Linné’s notion that these forms were due to hybridization. It is
extracted from Gmelin’s edition of the ‘Systema Nature,’ 1791, p. 931.
“ Tinarie proles hybrida, ejusdemque qualitatis et constans, radicibus
infinite sese multiplicans charactere fructificationis diversissima, corolla
regulari, quinque-corniculata, pentandra, ut genus proprium absolute
constitueret et distinctissimum, nisi fructus frequentissime abortiret.
Nature prodigium. Ita quidem a Linné. Verisimilior autem videtur
ea opinio, que peloriam pro peculiari degeneratione monstrosa floris
habet, in quam inclinare hoc genus (Linaria) pre aliis, similis a forma
deflexio in aliis speciebus, e.g. spurio Elatine, eymbalaria, observata,

ae Merk., ‘Goett. gel. Anz.,’ 1774, n. 121. Linck, ‘ Annal.
Nature.,’ i, p. 32.”

230 IRREGULAR PELORIA.

serve to show in what way the irregularity is brought
about. In Antirrhinum, Iinaria, &c., mtermediate
forms show very clearly that it is to the repetition of

Fig. 122.—Peloric flower of Calceolaric.

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! 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.

IRREGULAR PELORIA. 231

narrow orifice of the corolla. The colour at the base
of the tube inside is as in the perfect peloria; while
round the summit of the tube, in both cases, the 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 Cytisus Laburnwm, Clitoria
Ternatea, Trifoliwm repens, and other Papilonacee,
it is the ‘‘ standard,” the form of which is repeated.
In the case of peloric aconites’ 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.” 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,’ 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 Phalwnopsis equestris
sent to me by Mr. Wentworth Buller. Fig. 123 repre-
sents a flower of Avistolochia caudata 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. du genre Aconitum,’ p. 124.

2 Schlotterbec, ‘ "Act. Helv et.,’ t. ii, pl. i, Roeper. Balsam, p. 10, note.

3 Masters, “ Peloria, &e., Ophrys aranifera, *< Journ, Linn. Soc.,’ viii, p.
207.

Dae IRREGULAR PELORIA.

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

Fic. 123.—Two-lipped flower of Aristolochia caudata.

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 Kdinburgh, December 13th, 1860, four
abnormal flowers of the common Indian cress (Tvropeo-
lum majus), each presenting a supernumerary spur. On
these he remarked that ‘‘in Tropewolwm the posterior

IRREGULAR PELORIA. PAS bo.

part of the receptacle between the insertion of the
petals and that of the stamens is dilated so as to form
the spur which is so characteristic in the genus. The
position of the spur in a line with the posterior sepal
has led many botanists to consider it as a process of
that sepal, but the fact of its being situated within the
insertion of the petals 1s 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 im-
creased; thus, in Antirrhinwm majus the corolla, when
subjected to peloria, is very generally six-parted, and
has six stamens. Fusion of one or more flowers is also
a common accompaniment of peloria, as in Digitalis
purpurea, in which plant prolification often adds 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 Teucriwm campanu-
latwm, for instance, is very generally regular. In Calceo-
lavia it is the central terminal flower which is usually
peloriated ; on the other hand, in Iinaria and Antirr-

234: IRREGULAR PELORIA.

linwm 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 pomt, 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.’

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’ 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 foliage, 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.

Linné, 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, “ Mém. sur les Fumarieés 4 fl. irreg.,” ‘Ann. Se. Nat.,’ sér.

5, vol. ii, tab. xvii, p. 280.
2 «Bull. Soc. Bot. France,’ vol. v, 1858, p. 701.

IRREGULAR PELORIA. 935

seeds from the Linaria which reproduced the
anomaly when sown in rich soil. Baron Melicog
obtained similar results.’ Mr. Darwin’ raised sixteen
seedling plants of a peloric Antirrhinwm, 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 versd, 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. Asimilar 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 Innaria, 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 1s well known, accompany
the diversities in form.

To Mr. Darwin the author is indebted for the com-
munication of some flowers of Corydalis tuberosa (figs.
124, 125), provided with two spurs of nearly equal size.
To these flowers allusion is made in the work already
quoted® 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

Bull. Soc. Bot. France,’ vol. vi, 1859, p. 717.
* * Variation of Anim, and Plants,’ ii, p. 70.
3 Loe. cit., p. 59.

236 IRREGULAR PELORIA.

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. 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 imsect’s body. In
several closely allied genera, as in Dielytra, there are
two perfect nectaries ; the pistil is straight, and the
hood slips off on either side, according as the bee sucks
either nectary.” In the flowers of Corydalis, which
were provided with two perfect nectaries contaiing
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 Dielytra 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

IRREGULAR PELORIA. 237

deviations from the ordinary form of one plant repre-
sent the ordinary condition of another ; thus, the peloric
Calceolarias resemble the flowers of Fabiana, and De
Candolle,' comparing the peloric flowers of Scrophula-
riacee with those of Solanacew, concluded that the
former natural order was only an habitual alteration
from the type of the latter. Peloric flowers of Papi-
lionacee im this way are indistinguishable from those
of ftosacee. In like-manner we may trace an analogy
between the normal one-spurred Delphiniwnm and the
five-spurred columbine (Aquilegia), an analogy
strengthened by such a case as that of the five-
spurred flower of Delphiniwm elatum described by
Godron.” The Corydalis, before referred to, is another
illustration of the same fact, the structure being the
same as in Dielytra, &e.

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
Iinaria 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 peloriz, solo sterili plantate, degenerant
in Linariam,”’ (‘ Sp. Plant.,’ i, p. 254); but this opinion
is counterbalanced by that of others, while the frequent

1 *Théor. Elém.,’ ed. 2, p. 266.
? Cited in ‘ Bull. Soc. Bot. France,’ vol. xiii (Rev. Bibl.), p. 81.

238 IRREGULAR PELORIA.

existence of both forms on the same plant, at the same
time, seems to negative the supposition of any direct
effect from external circumstances.

The following are the plants in which irregular
peloria has been most often observed :

Aconitum Napellus.
Delphinium elatum !
Corydalis tuberosa.
*Viola odorata !
hirta.
Impatiens Balsamina.
Clitoria Ternatea.
Cytisus Laburnum!
Trifolium repens !
Lupinus polyphyllus !
*Gloxinia, var. cult. !
*Linaria vulgaris !
spuria.
Hlatine.
triphylla.
eruginea.

triornithophora.

pilosa.
chalepensis.
cymbalaria !
purpurea !
decumbens.
Pelisseriana.
origanifolia.
Digitalis orientalis.
: purpurea !
Calceolaria crenatiflora.
rugosa.
* var. cult. !
Chelone barbata.
*Antirrhinum majus !

Rhinanthus crista galli.

Pedicularis sylvatica.

Pedicularis euphrasioides.
Scrophularia aquatica !
Sesamum indicum,
Lamium.

Mentha.

Sideritis.

Nepeta diffusa.

Galeopsis Ladanum.

Tetrahit.

Galeobdolon luteum.

Teucrium campanulatum!

Plectranthus fruticosus.

Cleonia lusitanica.

Dracocephalum austriacum.

Phlomis fruticosa !

Vitex incisa.

Aristolochia, sp. !

Ophrys aranifera !

Orchis simia.
pyramidalis !
latifolia !
morio !
papilionacea.
mascula.
latiflora.
conopsea.

Habenaria bifolia.

Corallorhiza innata.

Aceras anthropophora.

Cattleya Mossi !

Phalnopsis 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).

Linwria.—Adanson, ‘Fam. Plant.,’ t.i, p. 110. Jussieu, ‘ Gen. Plant.,’
p. 120. Poiret, ‘Encycl. Method. Suppl.’ t. iu, Jaeger, ‘ Missbilld.
der Gewachs.,’ pp. 94, 97, and 313. Cassini, ‘Op. Phytol.’ t. ii, p. 331.
Ratzebourg, ‘ Animadv. ad pelor. spectand.,’ 1525. Turpin,‘ Ie. Veget.,’

IRREGULAR PELORIA. 239

tab. XX, TGs (Curtis, ‘Flor. Londin.,’ i, 118. Hopkirk, ‘ Flora Anom.,’
pl. vii, figs. 1, 2 2, 3. Haller, ‘ Act. Helvet.,’ 2, p. 25, t. iv. De Candolle,
‘Flore Franc.,’ t. ili, p. 583. Sowerby, “Engl. Bot.,” iv, 260, ed, Syme,
tab. 963. Chavannes, ‘Mon. Antirrhin.’ Delavaud, ‘Bull. Soe. Bot.
France,’ 1858, p. 689; id., 1860, p. 175. Heufler, ‘ Linnea,’ xvii, tab. ii.
Weber, ‘ Verhandl. des Nat. Hist. Vereins. t-d. Rh. Preuss.” 1850, tab. i,
figs. I-8. ‘Verh. Nat. Hist. Ver. Rh. Preus.,” 1849, vol. vi, p. 290,
tabs xii.—Antirrhinwm, Clos, ‘Mém. Acad. Toulous. ” vi, 1862. Cha-
vannes, ‘Mon. Antirrh., p. 62. Fresenius, ‘ Mus. Senkenb.,’ Vata! LVis
fig. 10. ‘Bot. Soc. Edinb.,’ 1851, July 10.—Calceolaria, Chamisso,
‘Linnea,’ t. vii, p. 206. Guillemin, ‘Archiv. Bot.,’ t. ii, p. 1 et 136.
Schlechtendal, ‘Linnea,’ xii, p. 686. Ernst Meyer, ‘ Linnea,’ xvi, 26,
tab. iii. Morren, ‘ Bull. Acad. Belg.,’ t. xv, n. 7, et t. xviii, P- 583.
‘Gard. Chron.,’ 1850, p. 389; ibid., ” 1866, p. 612.—Viala, Leers, ‘ Flor.
Herborn..,’ p. 145. De Candolle, ‘ Organ. Veget.,’ p. 519, pl. xlv.
Forbes, ‘Proc. Linn. Soc.,’ June 6, 1848, p. "399, rtdebeand: ‘ Bot.
fod > 1862, vol. xx, tab. viiii—Orchidacee, His, ‘Jourl. Phys., 65,
_ 24d, Wydler, ‘Arch. Bot.,’ t. ii, p. 310, tab. xvi. R. Brown, ‘ Obs.
Bean Orchid.,’ p. 698. A. Richard, ‘Mém. soc. Vhist. 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, ‘Mém. Acad. Sc. Toulous.,’ 5 ser., vol. iii. Caspary,
‘Schrift. K. Gesellsch. Koénigsberg,’ 1860, i, 59. Masters, ‘ Jourl.
Linn. Soc.,’ vol. viii, p. 208 (Ophrys, Pogonia). Duchartre, ‘ Bull. Soc.
Bot. Fr.,’ vol. vii, 1860, p. 26, Cattleya. Cramer, ‘ Bildungsabweich.’—
Limosella, Baillon, ‘ Adansonia,’ i, p. 305. (Flower normally irregular,
becoming regular “‘a force d'irregularité.”)—Chelone, Chamisso,
‘Linnea,’ vii, p. 206.—Clitoria, Bonavia, ‘Gard. Chron.,’ 1868, p.1013.
In this latter communication, published as this sheet is passing thr ough
the press, the author gives an interesting account of the transitional
stages between the ordinar 'y papilionaceous condition and the regular
form which is like that of a Rosaceous plant. The peloric form is stated
to be transmitted by seed.
For other references see Moq.-Tandon, ‘ El. Terat. caeaet p. 186.
Hallier, ‘ Phytopathol., p. 151.

PA Rod al
METAMORPHY.

Mucu 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 i 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 fiower, 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 imphes an
alteration in the organizing ace. 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

PHYLLODY. 2A]

elsewhere, but, under the present heading, are specially
included cases not of merely diminished or increased,
but of perverted development; the natural process is
here not necessarily checked or enhanced, but it is
changed. Hence, in the present work, the term
metamorphy is employed to distinguish cases where
the ordinary course of development has been perverted
or changed. As it is applied solely for 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.’

CHAPTER I.

PHYLLODY.

THis condition, wherein true leaves are substituted
for some other organs,” must be distinguished from
Virescence, q.v., 12 which the parts affected have
simply the green colour of leaves, without their form
or structure. The appearance of perfect leaves, in

! 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,
Linné, 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, :

16

242, PHYLLODY

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 mstance,
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

Fie. 126.—‘ Rose plantain,’ Plantago media var., spike contracted ;
bracts leafy.

OF THE BRACTS. 243

is not attributable to arrest of development or retro-
erade metamorphosis, but the reverse. The bracts of
some species of Plantago' are very subject to this change.
Thus, in the rose plantain of gardens, P. media (fig.
126), the bracts are leafy and the axis depressed or
not elongated, so that it is surmounted by a rosette of
small leafy organs. <A similar condition of the bracts,
unattended with arrest of growth in the axis, is common
in P. major (fig. 127) and in P. lanceolata (see p. 108).

Fic. 127.—Leaf-like bracts in Plantago major.

It also occurs in the bracts of Corydalis solida, Amorpha
fruticosa, Ajuga reptans, Parthenium inodorum, Cent-
aurea Jacea, in the involucral bracts of the dandelion, the

1 See Schlechtendal, ‘ Bot. Zeit.,’ vol. xv, 1857, p. 873; also Marchand,
* Adansonia,’ iv, p. 156.

244, PHYLLODY

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 allassumed the form, texture, and venation of leaves.

SS.

=S¢

SA
qv

Fie. 128.—Dahlia. Scales of receptacle leafy.

In Umnbellifere the substitution of leaves for involucral
bracts is not infrequent. It has been observed among
other plants in Angelica Razoulzii, Carum carui, Daucus
Carota, &e. The scales of the hop (Hiwmulus Iupulus)
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,”
of some willows,® &c. The bracts of some Huphor-
biacece, as H. pusilla, EH. Lathyris, EF. Cyparissias, have
been observed to undergo a similar alteration.‘

' For instances of similar changes in Composites, see De Candolle,
‘Prod., t. vi, p. 571, Centaurea Jacea phyllocephala. Clos, ‘ Ann. Se.
Nat.,’ ser. i, tom. xvi, 1851, p. 41. ‘Science Gossip,’ 1865, p. 104, &e.

2 Kickx, ‘ Bull. Acad. Belg.,’ t. xviii, part 2, p. 288.

3 Weber, ‘ Verhandl. Nat. Hist. Vereins. f. Preuss.,’ &c., 1860, p. 381.

* Weber, loc. cit. ;

OF THE CALYX. 245

Amongst monocotyledons an analogous change
occurs not unfrequently, as in some commelynaceous
plants, e.g. Tradescantia, in Musa, &ce.

The spathe of Arwi maculatum 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-lke spathe.' In Schenus cephalotes a similar
exaggerated development of the bracts is figured by
Rottboell.”

Phyllody in inflorescence of Conifers——'T'his demands pass-
ing notice by reason of the interest attaching to the
morphological construction of these plants. The
elongation of the axis which occurs im 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 Parlatore in Abies Brunoniana, and examples
may frequently be met with in the larch (Larizv europea),
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 Araucaria, as also in Podocarpee
and Cupressinee (Hichler).

Phyllody of the calyx——Sepals under ordinary circum-
stances are so like leaves, that it is not wonderful that

' Sauter, ‘ Flora v. Bot. Zeit.,’ 1831, p. 11.

2 *Descr. et Icon. Plant.,’ tab. 20.

* For references see p. 115; see also to Eichler, ‘ Excurs. Morpholog.
de format. flor. Gymnosperm.,’ in “‘ Mart. Flor. Brasil,” abstracted in
English in ‘ Natural History Review,’ April, 1864.

246 PHYLLODY

they are often replaced by those organs.’ A singular
instance of this has been mentioned as occurring in
Cakile maritima, wherein the sepals were found by
M. Fournier to be pinnatifid like the ordinary leaves
of the plant.” The sepals of Ranunculacee and Rosacee,
for example, fosa, Geum, are particularly liable to this
change.

Fic. 129.—Flower of rose, sepals replaced by five perfect leaves; axis
prolonged through the flower in the form of a leafy branch.

In a species of Geranium recently examined the
sepals presented themselves in the form of three-lobed
leaflets; so in fuchsias and in Hpilobium hirsutum the
sepals occasionally are not distinguishable from ordinary
leaves (fig. 130). In roses, the change in question is a
very frequent accompaniment of prolification (fig. 129).
In the peach also this replacement of the sepals is

Li Calyx tune plane non differt a foliis proxime ipsi precedentibus.”
Wolff, ‘Theor. Gener.,’ § 114. Linn., ‘ Proleps.,’ § 6. Goethe, ‘ Ver-
such.,’ §§ 31-38.

2 ‘Bull. Soc. Bot. France,’ vol. viii, 1861, p. 697.

OF THE CALYX. 947

sometimes carried to such an extent, that five perfect,
bistipulate leaves occur in the place of the calyx, but
when this is the case it usually happens that the
pistil is abortive.

Fie. 130.—Fuchsia, with one of the sepals leaf-like.

De Candolle’ figures a curious instance wherein the
pappus of Podospermum laciniatum was replaced by
five linear, foliaceous lobes. A similar change has
been noticed in other composites, as in Tragopogon
pratense. Engelmann mentions as subject to this
hypertrophy of the pappus, as it may be termed,
Scorzonera octangularis and Senecio vulgaris. Wigand
has observed a similar transformation in a species of
Centranthus (Valerianacee).

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-hke organs sometimes seen at the
apex of a cucumber would seem to support the view
that there was really a calyx-tube in Cucurbitacee
adherent to the carpels. It is also shown in the cut,

1 «Organ. Véget.,’ t.i1, p. 492, pl. xxxu, f. 6.

24S PHYLLODY

fig. 132, borrowed from the ‘ Gardeners’ Chronicle,’
1859, p. 654.

Fie. 152.—Leafy calyx of melon.

OF THE CALYX. 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
cases; 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.’

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 lamimar 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

! This distinction between laminar and vaginal venation is well seen
in cases like Mussaenda, Calycophyllum, or Dipterocarpus, 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—“ soudée
au calice et ayant dévorée, 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).

250 PHYLLODY

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.
incanus.
Sinapis arvensis.
Brassica oleracea!
Peltaria alliacea.

*Sisymbrium officinale.
CaryophyLlacez,! sp. pl.

Geranium, sp. !

*Puchsia, var. hort. !
Epilobium hirsutum !
Cucurbita Pepo !

* Rosa, var. hort. !
Potentilla nepalensis.
Fragaria sp.

Geum rivale.

Persica vulgaris.
Cerasus !
Pyrus Malus.
Daucus Carota.
Athamanta Cervaria.
*Trifolium repens !
Centranthus macrosiphon.
Tragopogon pratense.
orientale.
Scorzonera octangularis.
Hypocheris radicata,
*Senecio vulgaris !
Podospermum laciniatum.
Cirsium arvense.
Carduus heterophyllus.
tataricus.
Campanula, sp.
Convolvulus sepium.
* Primula officinalis, var. cult !
acaulis.
elatior.
Gentiana campestris.
*Petunia violacea!
Lycium europeum.
Laurus Sassafras.
Tulipa Gesneriana.
Convallaria maialis.
Colchicum autumnale! (vire-

Amygdalus communis. scent ?)

Consult also Turpin, ‘Atlas de Goethe,’ t. iv, f. 12, Zycium. Engel-
mann, ‘De Anthol.,’ § 35, p. 31. This author figures phyllodie 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, &e. ‘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.

1 In this order Agrostemma Githago offers an illustration of a normally
leafy calyx.

OF THE COROLLA. 251

Phyllody of the corolla——'T'he petals also are frequently
replaced by leaves, though in many of the recorded
instances the change has been one of colour only;
these latter are strictly cases of virescence. M. Seringe!
speaks of a flower of Peltaria alliacea in which the
calyx was petal-like, while the corolla was leafy as if
there had been transposition of the two organs, a very
rare, if not unparalleled, instance. In a flower of
Campanula Mediwm, 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 Verbascwm nigrum, and Lonicera Peri-
clymenum, 1 which, moreover, median prolification
generally coexists. In the case of Tropeolum 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 ves 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 Actea
spicata, as observed by Fresenius, the petals were
replaced by true petiolate, palminerved, lobed leaves,

' «Bull. Bot.,’ i, p. 6.

252, PHYLLODY

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 Tropaolum 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 lamine.
(See the remarks in the preceding section.)

Frondescence of the petals has been observed most
frequently in the following cases; some, perhaps, were
cases merely of virescence, q. v.; see also under
Chloranthy, Prolification.

Ranunculus repens !
Delphinium Ajacis.
crassicaule.
Aquilegia vulgaris.
Actza spicata.
*Brassica oleracea !
Diplotaxis muralis.
Hesperis matronalis.
Thlaspi bursa pastoris.
Sisymbrium tenuifolium.
Turritis glabra.
Raphanus sativus.
Peltaria alliacea.
Alyssum incanum.
Erysimum Barbarea.
officinale !

cheiranthoides.

Cheiranthus Cheiri.
*Dictamnus Fraxinella!
Lychnis sylvestris.
dioica!
Alsine media.
Cerastium vulgatum !
triviale.
Reseda lutea.
Phyteuma.
Malva sylvestris.
*Tropeolum majus !

Geranium, sp.!
Triumfetta, sp.!
Epiulobium hirsutum !
(nothera striata.
Rubus, sp.
*Rosa, var. cult. !
*Trifolium repens !
Spirea oblongifolia.
Amygdalus communis.
*Rosa!
Cerasus vulgaris !
Persica vulgaris !

' Potentilla nepalensis.

Geum rivale.
Daucus Carota!
Heracleum Sphondylium.
Torilis Anthriscus.
Eehinophora maritima.
Campanula rapunculoides.
glomerata.
Phyteuma spicatum.
Calendula officinalis.
Cirsium tricephalodes.
Senecio vulgaris.
Seabiosa columbaria.
agrestis.
Lonicera xylosteum.
Periclymenum.

OF THE STAMENS. 253
Gentiana Amarella. Antirrhinum majus !
Gilia glomeriflora. Stachys sylvatica.
*Symphytum officinale. *Anagallis phcenicea ?
Petunia violacea! Primula sinensis !
Verbascum, sp. Polemonium cceruleum.

See Moquin-Tandon, ‘El. Terat. Veg.,’ p. 203. Engelmann, ‘ De
Anthol.,’ § 38 et seg. ; tab. ii, figs. 8—14, Gilia ; tab. v, 23—26, Senecio ;
tab. v, f. 1—13, Torilis ; tab. iv, f. 3, Hrysimum. ‘Bull. Soc. Bot. Fr.,’
vol. ii, 1855, p. 479, Primula sinensis. Giraud, ‘ Edinb. Phil. Magazine,’
1839, Antirrhinum. Jaeger, ‘ Act. Acad. Ces. Nat. Cur.,’ vol. xiii, 2,
p. 1, tab. xli, Tropeolum. Bischoff, ‘Lehrbuch,’ 11, 2, p. 27, note,
Tropeolum. Fresenius, ‘Mus. Senkenb.,’ ii, 35, tab. 4, fig. 5, Actea. See
also succeeding paragraphs and sections in Chloranthy, Virescence, Xe.

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.’

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, i which, im addition to other
changes mentioned under prolification of the inflor-
escence, some of the stamens were partly frondescent,

1 Wolff’s original opinion was that the stamens were equivalent to so
many buds placed in the axil of the petals or sepals (see ‘ Theoria Gene-
rationis,’ 1759, § 114)—an opinion which more recently has received the
support of Agardh and Endlicher. Wolff himself, however, seems to
have abandoned his original notion, for in his memoir, “ De formatione
intestinorum precipue tum et de amnio spurio aliisque partibus embry-
onis gallinacei, nondum visis,” &c., in ‘Comm. Acad, Petrop.,’ xii, p. 403,
anno 1766, he considers the stamens as essentially leaves. See also
Linn. ‘ Prolepsis,’ § viii; Goethe, ‘ Metam.,’ § 46.

254: PHYLLODY

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-

Fie. 134.— Flower of a Petunia, opened to show the stamens partially
replaced by stalked leaves.

duplicate, and a third central one, half anther, half
leaflet.

In the case of frondescent flowers of Tropeolum
majus the stamens are usually absent or atrophied,
but in other instances the filament is present as usual,
representing the stalk of the leaf, and surmounted by
a small lamina, but this latter, in place of being nearly
flat, is pinched up in the centre from back to front,
and surmounted by a two-lobed anther, so that the
general appearance of the whole structure is that of
a central anther, supported at the base on each side by
two concave leaf-lobes, or it might be compared with
a three-lobed leaf, the terminal lobe represented by
the anther.

In Jatropha Pohliana, Mill. (Adenorophium lueu-
rians, Pohl.), a singular condition has been observed by
M. Miller (Argov.). In this flower the anther, in place

OF THE STAMENS. 955

of being represented by the flat blade of a single leaf,
had the appearance as if two such blades were present
and coherent one with the other by their midribs,
along their upper or inner surfaces, which were directed
towards the centre of the flower (fig. 136), thus resem-
bling the cases of adhesion of leaves by their surfaces
already referred to (p. 33). In other cases, in the same
plant, the anther appeared as if formed by two collateral

Fig. 135.—Phylloid anther of Fic. 136.—Leaf-like anther of
Jatropha, after Miller (Arg.). Jatropha Pohliana, after Miller.

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. 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.

Phyllody of the stamens has been most often observed
in the following plants :

' Miiller (Argoy.), in ‘ Mém. Soc. Phys. et d’Hist. Nat. Genev.,’ t. xvii.

256 PHYLLODY

Anemone nemorosa. Daucus Carota.
coronaria. Epilobium hirsutum !
Delphinium crassicaule. *Rosa, var. cult. !
Nymphea dentata. Lonicera Periclymenum.
Tropzolum majus ! Anagallis arvensis.
Dictamnus albus. Primula sinensis!
*Trifolium repens! Petunia, var. cult.
Torilis anthriscus. Jatropha Pohhana.
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’—This 1s 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 bemg here represented by two
small fohar laminze, 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’
relates having found in the neighbourhood of Mont-

eller a flower of a tulip the ovary of which was repre-
sented by true leaves, which bore on their margins the
ovules, and thus presented a striking analogy with
the carpels of those Stercuhas, like S. platanifolia,
which are foliaceous in texture and open very early in
the course of their development. A similar occur-

1“ Tf 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. 405, expresses precisely the same opinion as to the nature
of the seed-vessel.

2 «il. Terat. Veg.,’ p. 205.

OF THE CARPELS. 257

rence has also been frequently noticed in the Columbine
and also in Orucifere and Umbellifere. M. Germain de
St. Pierre mentions an instance wherein the carpels of
Salix Babylonica were converted into two leaves, pro-
vided with stipules. All the flowers of the catkins

Fia. 137.—Rose, in which the axial portion of the flower was elon-
gated and the carpels were more or less replaced by leaves.

17

258 PHYLLODY

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 Ranunculus and Anemone.

In proliferous roses, or in cases where the central axis
of the flower is prolonged, it frequently happens that the
pistils are more or less replaced by leaves. Fig. 137,
from a specimen of Dr. Bell Salter’s, given in the
‘Gardeners’ Chronicle,’ shows the passage, from below
upwards, of the ordinary carpels to perfect leaves ;
the so-called calyx-tube being completely deficient
and the ovaries entirely superior. Like most similar
specimens, this one bears out the notion that what is
called the calyx-tube in roses is really an expansion
and dilatation of the top of the flower-stalk.

Fic. 138.—Cucumber with leaf attached.

OF THE CARPELS. 259

Fig. 138, for which I am indebted to Mr. 8. J.
Salter, represents a very singular conformation in the
cucumber, described by that “eentleman in ‘Henfrey’s
Botanical ‘Gazette,’ i, p. 208, and considered by him to
be due to the foliaceous condition of one of the three
carpels of which the fruit is composed. The portion
near the peduncle was binary, while the distal extremity
of the fruit was ternary. he 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 Cucurbitacee 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 meght 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’ 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-
dition.

1 « Ann. Se. Nat.,’ 4th series, vol. ix, p. 209.

260 PHYLLODY

In the preceding instances the foliaceous condition
has pervaded the entire pistil, or at any rate the basal

Fic. 159.—Flower of Triumfetta, sp., carpels represented by five
leaves.

portion or ovary, and it may be noticed that the ovary
is thus shown to consist in some cases of the sheath
of the leaf, as in Aquilegia ; in other cases of the blade,
as in Cerasus, Daucus, &e.

There are cases, however, in which a part only
of the pistillary structure thus becomes foliaceous.
Linnezeus, ‘ Prolepsis,’ § 9, mentions some flowers of
Carduus heterophyllus and C. tataricus in which the
style had grown into two green leaflets, and in which
the calyx and corolla were also leaf-like. A very
singular instance is recorded by Baillon,’ 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 placentz ; 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-foholate leaves. The back of the leaf in these
cases 1s 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 Tropeolum majus, ‘ Bull. Soc. Bot. Fr.,’
1865; p, 411.

OF THE CARPELS. 261

flower. When this change occurs it is commonly
attended by an increased number of parts, as in the
trefoil just mentioned, or in the double cherry, where
usually two foliaceous carpels may be met with, and
sometimes more.

The change is also of interest when it affects such
orders as the Umbelliferw, which have their ovaries
inferior under ordinary circumstances ; but when these
organs assume a leafy condition they become superior
also, 7. 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,
Delphinium, &c., on the edges of the carpel or on the
surface, as in some flowers of Ranunculus repens and.
R. Ficavia. 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.’ 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 :

Peeonia officinalis. Nymphea dentata.
Ranunculus repens ! Sinapis arvensis !
*A quilegia vulgaris! Diplotaxis tenuifolia.
Delphinium elatum. *Brassica oleracea !
crassicaule. *Sisymbrium officinale !
Ajacis. Dianthus, sp.
amenum. Reseda Phyteuma.

i Planchon et Marés, ‘ Ann. Sc. Nat.,’ ser. 5, vol. vi, 1866, p. 228, tab.
X11.

262

Triumfetta, sp. !
Lychnis dioica.
Cerastium, sp.!
*Dictamnus Fraxinella !
Cerasus avium.
vulgaris !
*Rosa, var. cult.! ~
*Daucus Carota!
Heracleum, sp.
Epilobium hirsutum !
‘Lathyrus latifolius.
*“Trifolium repens !
hybridum.

PHYLLODY

Melilotus, sp.
Medicago, sp.
Lonicera Periclymenum.
Carduus heterophyllus.
tataricus.
Scrophularia aquatica.
Symphytum officinale.
Anchusa ochroleuca.
paniculata,
*Primula sinensis !
Salix babylonica.
Hyacinthus, sp.
Tulipa, sp.

Some of the above are probably cases of mere
virescence rather than of phyllody. For further illus-
trations, references to authorities, d&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-
seribes 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 Primulacee, 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

OF THE OVULES. 963

ovules of Primula sinensis, applied the term lepyro-
phylly (Aerupdy, a scale) to the foliaceous condition of
the testa in this plant. Unger’ describes a series of
malformations in Primula sinensis, consisting chiefly of
reversions of the part of the flower to leaves. ‘The
carpels were entirely absent in this case, and the place
of the free central placenta was occupied by a circle of
leaves, sometimes bearing imperfect ovules on their
edges. An instance of a similar kind has been de-
scribed by A. de Candolle.°

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
Cortusa 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. Dr. Marchand* mentions
similar changes in Anagallis arvensis and Lonicera Peri-
clymenum.

Cramer’ figures ovules of Primula sinensis in the
form of stalked leaves, often becoming infolded at the
margins, and giving origin to a small nucleus on their
inner surface.

M. Tassi® records an instance in Symphytwin officinale
wherein the ovules were replaced by two small lnear
leaves arising entirely from the axis, and not from the
carpels.

In most of the foregomg illustrations the foliar portion
of the ovule must have been independent of the carpel ;
this independence is less manifest, though probably as

| * Act. Acad. Nat. Cur.,’ 22, 11, 1850, p. 543, t. v, vi.
2 * Neue Denkschrift der allg. Schweiz. Gesellsch.,’ band v, p. 9, tab. 3, 4.
3 * Ann. Se. Nat.’ 2 ser., vol. i, p. 308, pl. ix, ¢
4 * Adansonia,’ vol. iv, pp. 159, 171.

Bae Bildungsabweichungen,’ &e., tab. iv, figs. 1, 2, 21, 28, 29, &c.

® «Bull. Soc, Bot. Fr ance,’ vill, p. 395.

264 PHYLLODY

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.

Fia. 140.—Sinapis, replum and ovules; the dotted line shows the
position of the carpels.

Grifith, in alluding to a similar case in Sinapis,}
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 fohaceous 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 «Notule,’ p. 125, atlas, pl. xxxv; and ‘Journals of Travels,’ 1847,
p. 475, Lonicera.

OF THE OVULES. 265

In a species of T'riwmfetta (see p. 260), of which I
examined dried specimens, the
ovary was open and partly folia-
ceous; it bore on its imfolded
margins ten erect leaflets, repre-
senting so many ovules; each
leaflet was conduplicate, the back
being turned towards the pla-
centa.

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,'
from an examination of some
monstrous flowers of Drosera in-
termedia, was led to the inference yg, 141 —Leafy ovules,
that the ovules are analogous to &c., Trifolium repens.
hairs on the margins of the
leaves. This acute botanist was

Fie. 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.

266 PHYLLODY .

cup formed by the confluence of four glanduliferous
hairs and the concave leaf and the perfect ovule.
Brongniart’ records ovules of Delphiniwm elatwm
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. Clos? mentions a similar in-
stance in Aquilegia Skinneri ; and another is figured in
Lindley’s ‘ Hlements of Botany,’ p. 88, f. 180.
Cramer,’ 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

¥ie@. 143.—Portion of an open foliaceous carpel of Delphiniwm, with
ovules on the lobules.

' «Comptes Rendus,’ vol. xviii, March 25th, 1864, and ‘ Ann. Se. Nat.,’
3 ser., vol. ii, p. 32.

2 «Mém. Acad. Sc. Toulous.,’ ser. 5, vol. iii.

8 *Bildungsabweich, Pflanz. Famil.,’ p. 89, tab. xi.

OF THE OVULES. 267

from the modified lobes of the carpellary leaf. Figs.
143—145, copied from Cramer, show how the nucleus
of the ovule is formed as a new growth from the
surface of the lobes of the leaf in Delphiniwm elatum.

Fic. 144.—Section through marginal lobe of carpel (Delphiniwm), show-
ing the nucleus (7).

Fig. 145.—Section through marginal lobe of carpel, showing nucleus
and tegument (Delphiniwm).

One of the most singular instances of ovular mal-
formation on record is that cited by the Rey. 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

268 PHYLLODY

of the former, so that many intermediate stages could
be traced between the ordinary ovule and the ovary

Ap 2.

Fie. 146.—1. Placenta of Dianthus, bearing ovules and carpels.
2. One of the ovaries separated.

Fia. 147.—Ovules of Dianthus 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 c, d, e. Could such a change occur in the
animal kingdom, there would be the unfertilised ovum

OF THE OVULES. 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 was
not developed, and sufficient evidence has been above
given as to the fohar nature of the primine, 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 foliaceous
ovules there is never more than one coat, the secondine
and other integuments do not make their appearance
in these cases, and that very generally the change in
question accompanies a similar foliaceous condition in
the carpel, the margins of which are more or less dis-
united.

Prof. A. Braun remarks that up to this date no such
change has been observed in the ovules of Monocoty-
ledons.

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 1s 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. Wydler has observed a similar
occurrence in ovules of Alliaria officinalis, and Schimper
has described and figured specimens of Nigella dama-
scena 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 oyule!

Fig. 148 shows a floret of a species of Gaillardia,
im which the ovule was replaced by a leafy shoot which
had made its way through a chink in the ovary. In

-

270 PHYLLODY

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.

Fie. 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 fohar 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 Simapis and Lonicera, while Caspary’s conclu-
sions from the foliaceous ovules of Trifolium repens are
somewhat similar. The latter observer considers that
the funiculus, with the integuments, is the equivalent
of aleaflet, 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.

OF THE OVULES. 271

M. van Tieghem’s conclusion’ from the examination
of flowers of Tropeolum majus, in which the ovules
were replaced by perfect peltate leaves, is that the
ovules are foliar productions springing, not directly
from a prolonged floral axis, as in Primulacee, but
from branches of the axis arising from the axils of the
carpellary leaves.

Phyllody of the ovules has been met with most often
in the following species :

*Aquilegia vuigaris ! Pastinaca sativa.
Skinneri. Torilis anthriscus.
Delphinium crassicaule. Thysselinum palustre.
elatum. Epilobium palustre.
dictyocarpum. Rosa, sp.
Ajacis. agaria alpina.
Nigella damascena. *Trifolium repens !
Adonis autumnalis. Medicago maculata.
Cheiranthus Cheiri! Desmodium canadense.
Nasturtium, sp. Melilotus macrorhiza.
Sisymbrium officinale! Lonicera, sp.
Brassica napus ! Gaillardia !

* oleracea ! Crepis, sp.

*Alharia officinalis ! Phyteuma odorata.
Sinapis arvensis ! Symphytum Zeyheri.
Turritis, sp. ¥ officinale.
Thlaspi arvense. Stachys sylvatica.
Erucastrum Pollichii. Anagallis arvensis.
Stellaria media. pheenicea.

*Reseda lutea. Lysimachia ephemerum.
Drosera intermedia. *Primula sinensis !
Agrostemma Githago. Auricula.
Stellaria media. prenitens.
Triumfetta, sp. ! Gilia glomeruliflora.
Tropzolum majus ! Rumex arifolius.
Dictamnus albus. scutatus.

Fraxinella ! Salix caprea.

Carum carui.

The following list of publications relating to ovular
malformations is copied from A. Braun, ‘ Ueber Polyem-
bryonie und Keimung von Czlobogyne’ (Appendix),* to
which are also added some others not alluded to by

1 «Bull. Soc. Bot. Fr.,’ 1865, p. 411. :
* Translated in ‘ Ann. Se. Nat.,’ 4th series, t. xiv, p. 24.

20S PHYLLODY

that author and not specially referred to in the pre-
ceding pages :

Jaeger, ‘ Missbilld. d. Gewichse,’ p. 78, 79, f. 47. Reper, ‘ Enum.
Euphorb.,’ 1824, p. 45, Delphinium.—Schimper, ‘ Flora,’ 1829, pp. 437-8,
et ‘Mag. fur Pharmacie de Geiger,’ 1829-30, pl. iv—vi, text wanting,
Primula, Reseda, Cheiranthus—Engelmann, ‘De Antholysi,’ 1832.—
Valentin, ‘ Act. Acad. Nat. Cur.,’ 1839, p. 225, Lysimachia.— Unger. ‘ Act.
Acad. Nat. Cur.,’ xxii, 11, 1850, p. 543, t. 5 B, Primula.— Flora (B. Z.)’,
1842, p. 369, t. ii, Trifolivm—Brongniart, ‘ Ann. Sc. Nat.,’ 1834, ii, p. 308 ;
also ‘ Archives Mus. d’Hist. Nat.,’ 1844, t. iv, p. 43, pl. iv, v, Primula.—
Reissek, ‘ Linnea,’ xvii, 1843, Alliaria.—Wydler, ‘ Denkshrift. d. Regensb.
Bot. Gesell.” 1855, iv, s. 77, t. vii, Alliaria—Wigand, ‘ Grundlegung
der Pflanzen Teratol.,’ 1850, p. 39, Turritis—Wigand, ‘ Bot. Untersuch-
ungen, 1853, p. 23, Rosa, Turritis, Crepis—Germain de St. Pierre,
‘LInstitut, 1853, n. 1051, p. 351—Rossmann, “Entwicklung der
Eiknospen aus dem Fruchtblatte,” &c., ‘ Flora,’ 1855, pp. 647 and 705.—
Dareste, ‘Ann. Se. Nat., 1842, p. 220, Delphinium.—Fresenius, ‘ Mus.
Senkenb.,’ 1, p. 39, t. iv, £. 9, Primula.—Schultz, ‘ Flora o. d. Bot. Zeit.,’
1834, xvul, p. 121, Nasturtium.—Seringe and Heyland, ‘ Bull. Bot.,’ 1—7,
Diplotaxis— Clos, ‘Mem. Acad. Toulouse,’ vi, 1862, Delphinium.—Morren,
C., ‘Bull. Acad. Belg.,’ xix, part ii, p. 519, Primula—Caspary, ‘Schrift.
d. Physik. @k. Gesell. zu Konigsberg, band ii, p. 51, tabs. U1, i.
Fleischer, ‘ Ueber Missbildungen Verschiedener Cultur Pflanzen.,’ &c.,
Esslingen, 1862. Cramer, ‘ Bildungsabweich,’ p. 68, &. &c., Trifolimm—
Moquin-Tandon, ‘El. Terat. Veg.,’ p. 206, Cortusa.—Guillard, ‘ Bull.
Soc. Bot. Fr.,’ 1857, vol. iv, p. 761, Stellaria.—Moelkenboer, ‘ Tijdschrift
v. Natuurl. Geschied.,’ 1843, p. 355, t. vi, vii, Primula.—Van Tieghem,
‘Bull. Soc. Bot. Fr.,’ 1865, p. 411, Tropeolum.

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 Leguminose; the scales of some
begonias ; the tendrils of Bignonia, Cobea, &e.

The presence of two small green lamine on the
outer side of the two posterior stamens in Antirrhinum
majus has also been met with. The adventitious
organs appeared as if they were developments from
the thalamus—a kind of foliaceous disc, in fact.

CHLORANTHY. 273

Chloranthy—The term phyllomorphy is applied to the
individual parts of the flower which assume the form
and appearance of leaves. By chloranthy it is to be
understood that all, or the great majority of the organs
of the flower assume these conditions.’ 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.

ah a
WV hi Wi Zz

\ Vez zea
Y=

jj

if |

Fic. 149.—Leafy petal of Epi- Fie. 150.—Chloranthy, &e. Epi-
lobiwm. lobiwm hirsutum.

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 meant
by that term, or whether they were cases of mere
virescence (green colour, without other perceptible
change), or of prolification (formation of adventitious
buds).. It is, therefore, quite possible that some of

1 The calyx is not unfrequently excepted.

18

274. PHYLLODY.

the instances to be now mentioned were not strictly
cases of chloranthy.

Seringe’ has described a malformation in Diplotavis
tenuifolia in which all the floral organs were replaced
by sixteen distinct leaflets which had preserved their
proper relative position. The Crucifere, of which
family the last-named plant is a member, are particu-
larly lable to this malfcrmation, as also are the
Rosacee, as will be seen from the following illustrations.
Roses indeed often exhibit alterations of this kind as
the commencement of prolification. There is also in
cultivation a rose’ 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
“Organogénie,” has shown that at a certain period
of the development of the ordinary rose flower the
ovary contains two collateral ovules, of which one

Fic. 151.—a. Open leafy carpel of ‘green rose,” with two deformed
ovules. 0b. Ovule separate. c. Primine removed. d. Secondine and
nucleus, with the bulbous end that pr eels through the micropyle.

1 * Bull. Bot.,’ t. i, p. 6.
2 oe ‘Theor. Horticult.,’ ed. 2, p. 84, f. 17.

CHLORANTHY. 275

becomes in process of time suppressed.’ Gewm coccinewm
has been found by Wigand with its flowers in this con-
dition.”

Lindley* figures a very interesting illustration in
Potentilla nepalensis, in which some of the flowers
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 Alpine strawberry is also
described as occasionally subject to this transformation.
In these flowers the calyx remains normal, while all
the other parts of the flower, even to the coating of
the ovule, assume a leaf-like condition.*

1 Gris, ‘ Bull. Soc. Bot. Fr.,’ 1858, vol. v, p. 261, and ‘Ann. Se. Nat.,’
ser. 4, vol. ix, p. 80. Planchon, ‘ Flore des Serres,’ vol. i, 1856, p. 129.

2 ‘Flora,’ 1856, p. 711.

3 ‘Theory of Horticult.,’ ed. 2, p. 90, f. 25.

* As considerable interest attaches to the “Plymouth strawberry,”
and very little is known of it in this country, or on the continent, the
author gladly avails himself of this opportunity of inserting an account
of it, for which he is indebted to the kindness of Dr. Robert Hogg.—
The Plymouth Strawberry (Fragaria vesca fructu hispido) is a sort of
botanical Dodo upon which many have written, and which few have
seen. Many years have elapsed since it was first discovered; and
although a century and a half have passed since there was any évidence
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.

Tt 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 Plimonth, whose daughter
had gathered and set the roots in her garden, in stead of the common
strawberry ; but she, finding the fruit not to answer her expectation,
intended to throw it away; which labour he spared her in taking it and
bestowing it among the louers of such varieties, in whose garden it is yet
preserved.” Doubtless one of those “ lovers” was his friend John 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 1 would not let this discourse passe without
giuing you the knowledge of it. It isin 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 like
ynto a double ruffe, in the midst whereof standeth the fruit, which, when
it is ripe, sheweth to be soft and somewhat reddish, hike vnto a straw-
berry, but with many small harmlesse prickles on them which may be
eaten and chewed in the mouth without any maner of offence and is
somewhat pleasant as a strawberry; it is no great bearer, but those

276 PHYLLODY.

Among Leguminose a partial leafy condition (frond-
escence), or a more complete degree of the same change,
(chloranthy) is not infrequent, particularly in Trifoliwm
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,’ 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 ordimary 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, green, with no trace of ovules. Sometimes a
second carpellary leaf, similar to the first, is formed; in other cases the
central axis of the flower is occasionally prolonged into a head of young
flowers—median prolification. In some few instances the calyx is not at
all altered, but the carpellary leaf is trifoliolate, or even quinquefoliolate,

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-
brigiz 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

766, M. Duchesne informed the world of the generosity of “M.
Monti, Docteur de Philosophie et de Médecine a Boulogne en Italie,”
who divided with him a dried specimen taken from his own herbarium,
“Ce présent prétieux m’éte toute incertitude sur la nature de ce Fraisier
et sur ses caracteres monstrueux. I] paroit ne pas avoir aujourd’hui
plus @’existence.”

1 «Bull. Soc. Bot. France,’ 1856, vol. ii, p. 477.

CHLORANTHY. 277

the corolla being then absent. The heads of flowers in this first form
have the aspect of little tufts of leaves.

2. Each of the teeth of the calyx is represented by a long stalk, 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
separate; the tube of the calyx is not deformed and seems to be formed
of the petioles of the sepals united by their stipules. In this second
class of cases the corolla is papilionaceous, the filaments free, the 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. foliaceous,
dentate primine only. In one case the carpel was closed above, gaping
below, where it gave origin to several leaflets, the lower ones oval,
dentate, like ordinary leaflets, the upper ones merely lanceolate, leafy
lobes, representing the primine reduced to afoliaceous 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,
arrested in its development and scarcely exceeding the tube of the
calyx within which it is crumpled up; stamens but little changed;
carpellary leaf on a short stalk, not exceeding the calyx tube, but the
ovarian porticn very long, and provided with abortive ovules.

These three groups will be found to include most of the forms under
which frondescence of the clover blossoms occurs, but there are, of course,
intermediate forms not readily to be grouped under either of the above
heads. Such are the cases brought under the notice of the British
Association at Birmingham in 1849 by Mr. R. Austen, in some of which
the petals and stamens even were represented by leaves.

Although, on the whole, chloranthy is most frequent
in the families already alluded to, yet it is by no means
confined to them, as the examples now to be given
amply show. Specimens of Nymphwa Lotus have been
seen in which all the parts of the flower, even to the
stigmas, were leafy, while the ovules were entirely
wanting.

Planchon' figures and describes a flower of Drosera
intermedia that had passed into a chloranthic condition,
excepting the calyx, which was unchanged; the petals,
like the valves of the ovary, were provided with sti-
pules, and were circimnate in vernation.

1 “Ann. Sc. Nat.,’ 3 ser., vol. ix, p. 86, tabs. v, vi.

278 PHYLLODY.

M. A. Viaud-Grand-Marais’ 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 in*Dictamnus Fravinella.

M. Germain de Saint Pierre has also recorded the
following deviations in the flowers of Ruwmew arifolius
and f. scutatus; m these specimens the calyx was
normal, the petals large, foliaceous, shaped like the
stem-leaves, the stamens were absent, the three carpels
fused into a triangular leafy pod, as long again as the
perianth, the stigmas normal or wanting, the ovule
represented by a thick funicle, terminated by a folia-
ceous appendage analogous to the primine.”

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 fi witans, the spikelet of which, as observed
by Wigand,° consisted below of the or dinary unchanged
elumes, but the remaining paleze as well as the lodicles
and stamens were represented by ligulate leaves. The
plant, it is stated, was affected by a parasitic fungus.
On the other hand, General Munro, in his valuable
monograph of the Bambusacee,' refers to an illustration
in which “the lowest glumes generally, and the lowest
palez occasionally, had the appearance of miniature
leaves, with vagine, heules and cilia, enveloping, how-
ever, perfect fertile spicule; as progress is made to-
wards the top of the spike, the leule 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

' * Bull. Soc. Bot. France,’ vol. viii, 1861, p. 695.
2 Thid., vol. iii, 1856, p. A755,

. ‘Flora,’ 1856, p. 712.

Se Trans. Linn. Soc.,’ vol. xxvi, p. 37.

CHLORANTHY. 279

remarks with justice that the position of the flowers
on the axis is of importance with reference to the
existence of chloranthy. Terminal flowers are more
subject to it than lateral ones, and if the latter, by
accident, become terminal, they seem peculiarly liable
to assume a foliaceous condition. Kirschleger says,
that im fiubus 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.’

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’ gives an instance
in Stellaria media where the condition appeared to be
due to the attacks of an insect Thrips fasciata. Still
more commonly it arises from the attacks of parasitic
fungi, e.g. Uredo candida, in Crucifers, Ke.

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 speciés 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,’ 1862, vol. ix, p. 36, tab. i, and also p. 291.
? Ibid., 1857, vol. iv, p. 761.

280 PHYLLODY.

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 m
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
inflorescence.

Selinum caruifolium.
Epilobium hirsutum !
Begonia fuchsioides.

Aquilegia vulgaris.
Chelidonium majus.
Corydalis aurea.

Nymphea Lotus!
*Brassica oleracea!
Bunias.
Hesperis matronalis.
*Sinapis arvensis !
Sisymbrium officinale.
Erucastrum canariense.
Diplotaxis tenuifolia.
Lychnis dioica!
Cerastium glomeratum !
triviale.
Stellaria media.
Poterium polygamum.
Torilis anthriscus.
Seseli, sp.

Gomphia, sp.
Scabiosa Columbaria.
Dipsacus fullonum.
Matricaria Parthenium.
Calendula officinalis.
Campanula pyramidalis.
Reseda odorata !
Vitis vinifera.
Dictamnus Fraxinella!
Triumfetta, sp. !
*Tropeolum majus!
Rhamnus Frangula.
*Trifolium repens !
Lupinus, sp.
Rosa diversifolia !

METAMORPHY 281

Potentilla nepalensis. Anagallis arvensis.
argentea. Webbiana,
Fragaria vesca ! Nicotiana rustica.
Geum rivale. Anchusa ochroleuca.
Rubus fruticosus. Myosotis cxspitosa.
cesius. Stachys sylvatica.
Saxifraga foliosa. Gilia capitata.
Verbascum phlomoides. Euphorbia segetalis.
Scrophularia nodosa. Rumex arifolius.
aquatica ! scutatus.
*Primula sinensis! Juncus lampocarpus.
Lysimachia Ephemerum. 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.

t. i, p. 308. A. P. and Alph. De Candolle in ‘Neue Denkschrift.’
Morren, C., ‘Bull. Acad. Roy. Belg.,’ xix, part 2, p. 539. Molkenboer,
‘Tijdschr. voor Natuurl. Geschied., 1843, p. 355, tabs. vi, vii. Marchand,
‘Adansonia, iv, p. 167 and p. 159, Anagallis, p. 171, Lonicera, p. 83,
Juncus. For other plants see Fresenius, ‘Mus. Senk.,’ 2, p. 35, &e.
Norman, ‘Ann. Sc. Nat.,’ ser. 4, 1858, vol. ix, p. 220. Christ, ‘ Flora’
(B. Z.) 1867, p. 376, tabs. v, vi, Stachys. Cramer, ‘ Bildungsabweich.,’
p. 26, &c. Baillon, ‘ Adansonia,’ ui, p. 300. Moquin-Tandon, ‘ El. Ter.
Veg.,’ p. 230. Schauer’s translation, p. 220. Haller, ‘ Phytopathologie,’
p. 160.

CHAPTER II.

METAMORPHY OF THE FLORAL ORGANS.

Onze 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,

282 METAMORPHY.

-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
Fi¢.152.—Flower Question there are neither stamens
of St. Valéry apple, nor petals, unless the second or inner
with sepaloid Pe- row of sepals be considered as sepa-

loid petals (fig. 152).

M. Alph. de Candolle’ 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 1s much less common than the
converse one of calycanthemy. Many of the recorded
instances of so-called metamorphosis of the parts of

1 ‘Neue Denkschrift. Schweiz. Gesellsch.,’ band v, p. 9.

PPTALODY. 283

the flower to sepals have occurred in monocotyledonous
plants, or others in which the calyx and corolla are of
the same colour, and constitute what is frequently
termed the perianth; and as this is usually brightly
coloured (not green) it is more convenient to group
the metamorphoses in question under the general term
Petalody, which thus includes all those cases in which
the organs of the flower appear in the form of coloured
petal-like organs, whether they be true petals or 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
colour.

Petalody of the calyx—Calycanthemy.— As with the bracts,
so the calyx in certain instances is naturally coloured,
as in Delphiniwm, Tropeolwm, and others. In Mus-
senda, Calycophyllum, Usteria, &c., one or more of the
calyx lobes become enlarged normally. Considered
teratologically, petaloid coloration of the sepals is
either general or partial; in the latter case the nerves
retain their green colour longest. There is in 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
ereat advantage in a horticultural pomt of view.
Morren' 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

' «Bull. Acad. Belg.,’ xix, part 2, p. 93.

284. METAMORPHY.

proper to the corolla is developed in the tube of the

Fie. 153.—Flower of Mimulus, with petaloid calyx.

calyx, the edges of the limb remaining green, the middle
of the imb being purple (Primula tricolor).

Under this head may be mentioned the occurrence of
tubular sepals in place of the ordinary flat ones in
Helleborus olympicus ; only two of the sepals were thus
affected im 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 :—Campanula persici-
folia, Anagallis arvensis, Gloxinia, Syringa persica,’
Calceolaria, Sc. Jc. In the last-named plant one or
more of the lobes of the calyx may frequently be seen
replaced by a slipper-lke petal.

or

1 Schlechtendal, ‘ Linnza,’ ix, p. 757.

PETALODY. 285

Among polysepalous plants petaloid sepals have
been observed in Ranunculus awricomus, Rubus cesius,
§c. Fleischer also describes a case of this kind in
Carum carui.'

It will be seen from the above that in the majority
of cases there is no real metamorphosis or substitution
of petal for calyx, but simply an alteration in colour ;
nevertheless, a charge in form may accompany a
change of colour: this happens especially if there has
been any displacement of organs. Thus, if, inan 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 versd.

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)’
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 Nymphea, Canna, and in some other plants,
such a transition occurs normally. Petalody of the
stamens may occur either without material change in
the flower or it may exist in combination or in 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.

1 Misbilld., ‘Cult. Gewachs.,’ p. 32.
2 Linn., ‘ Phil. Botan.,’ § 120.

286 METAMORPHY.

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 Delphinuwm.

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 Hrica Tetraliv 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 Jaliwm
Martagon, and there is in cultivation a variety of the
white lily, Liliwm candidwm, 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 tulips there is likewise a replacement
of stamens by coloured segments of the perianth, but
this happens generally in connection with an imerease
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 m this case was
transformed into several small leaves. Similar appear-
ances have been observed in Iris, Hyacinths, Nar-

PETALODY. 287

cissus, Colchicum, and Crocus. M. Fournier’ describes
a flower of Narcissus Tazetta from within the normal
perianth of which sprang a second one, equally pro-
vided with a cup and occupying the space usually filled
by the stamens. Flowers of Narcissus poeticus may
also be met with in which the stamens are replaced by
six distinct segments exactly resembling those of the
perianth in miniature.’

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.’ For instance, in Solanwm

Fic. 154.—Double columbine, Aquilegia—petalody of the filament.

1 «Bull. Soc. Bot. France,’ 1859, vol. vi, Pp. 199.

2 Seeman’s ‘Journal of Botany,’ vol. iii, p. 105; also Morren, ‘ Bull.
Acad. Belg.,’ vol. xx, part 2, p. 264.

3 Shige ioc ‘Bull. Belg.,’ xvill, p. 503.

288 METAMORPHY.

tuberosum, S. Duleamara, in Anagallis, in Fuchsia,
and some other plants, the anther-lobes themselves
become petaloid, while the filament remains un-
changed.

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’ observes that in the Ranunculacee 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-lke. Where the flower naturally
contains a large number of stamens, as in Mallows,
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 Alla-
manda cathartica, Jasminum grandiflorum, and many
other flowers with few stamens. A similar change im
the anther and connective takes place more frequently
in flowers where the number of stamens is smaller,
but there are of course numerous exceptions to this
rule.

In those cases where there is more than one row of
stamens, the outermost are most liable to this change :
thus in Sazifraga decipiens, as shown by Ch. Morren,”
the outer series of stamens—those opposite to the
sepals—become first affected, and, at a more advanced
stage, the inner row also ; and this is the case in most

1 «Organ. Vég.,’ t. i, p. 515.
2 «Bull. Acad. Roy. Belg., tome xvii; and Lobelia, p. 65.

PETALODY. 289

flowers that have their stamens in two rows. Occa-
sionally it happens that an outer series of stamens is
abortive, or wholly suppressed, while the inner row
becomes petalodic ; this was the case in some flowers
of Lilium auwratwin 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
jpetals, 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. Mogeridge.

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-

290 METAMORPHY.

tions being petaloid and continuous with the dilated
filament. Not unfrequently these semi-petaloid sta-

Fic. 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.’ (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 Pterandra,
and is common in some double fuchsias. In this
latter instance there is but a single wing, and the
nature of the case is obvious.

Double flowers of Orchidacew generally arise from
petalification of the filaments, with or without other
coincident changes. What makes double flowers in
this order the more interesting is the development, in
a petaloid condition, of some or all of those stamens
which under ordinary circumstances are wholly sup-
pressed, go that the morphological structure of the
flower, at first a matter of theory, becomes actually

1 Masters, “On Double Flowers,” ‘Rep. Internat. Bot. Congress,’
London, 1866, p. 127.

PETALODY. 29]

realised. Fig. 156 is a diagram showing the presence
of two additional labella within the ordinary one in
a species of Catasetwin, and repre-

senting two petaloid stamens, a
outer staminal whorl, of which

there is usually but a single

representative (see Peloria, Mul-

tiplication, Prolification). In =

some of these double orchids it is, Fxg. 156 —Diagram of
however, necessary not to con- flower of Catasetwm, with
found a petaloid condition of the *¥° #>ella.

existing column with the development of usually sup-
pressed stamens in a petaloid form. Thus, in Lycaste
Skinneri the column is frequently provided with two
petal-hke wings, which might readily be supposed to
be two stamens of the inner whorl adherent to the
eolumn ; 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.*

Some forms of duplicate or hose in hose corollas are
apparently due, not so much to the formation of a
second corolla within the first, as to the presence of an
inner series of petal-like stamens, which, by their
cohesion, form a second pseudo-corolla within the first.
The staminal nature of this pseudo-corolla is inferred
from the occasional presence of anthers on it.” In
Datura fastuosa, as well as in Gloxinia, a pseudo-
corolla of this kind sometimes occurs with the addition
of a series of petaloid stamens attached to its outer
surface.*

When the petalody specially affects the anther-lobes,
as in Arbutus, Petunia, Fuchsia, §c., the venation of
the petal-like portion is very frequently laminar, thus

thus evidently completing the f° \

! See also C. Morren, “Sur les vraies fleurs doubles chez les Orchi-
dées,” ‘ Bull. Acad. Roy. Belg.,’ vol. xix, part 11, 1852, p. 171.

2 ©. Morren, ‘Bull. Acad. Belg.,’ vol. xx, 1853, part ii, p. 284
(Syringa).

3 * Rep. Bot. Congress,’ London, 1866, p. 135, t. vii, f. 14.

299, METAMORPHY.

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-lhke 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 Although it is generally admitted that the filament of the stamen
corresponds to the stalk of the leaf, and the anther to the leaf-blade, yet
there are some points on which uncertainty still rests. One of these is
as to the sutures of the anther. Do these chinks through which the
pollen escapes correspond (as would at first sight seem probable) to the
margins of the antheral leaf, or do they answer to the lines that separate
the two pollen-cavities on each half of the anther one from the other ?
Professor Oliver, ‘ Trans. Linn. Soc.,’ vol. xxiii, 1862, p. 423, in alluding to
the views held by others on this subject, concludes, from an examination
of some geranium flowers in which the stamens were more or less petaloid,
that Bischoff’s notion as to the sutures of the anther is correct, viz.,
that they are the equivalents of the septa of untransformed tissue
between the pollen-sacs. Some double fuchsias (‘ Gard. Chron.,’ 1863,
p. 989) add confirmation to this opinion. In these flowers the petals
were present as usual, but the stamens were more or less petaloid, the 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
line separating the polliniferous from the petaloid portion of the anther.

: This view is also borne out by the double-flowered
Arbutus Unedo, and also by what occurs in some
double violets, wherein the anther exists in the
guise of a broad lancet-shaped expansion, from
the surface of which project four plates (fig. 157),
representing apparently the walls of the pollen-
sacs, but destitute of pollen; the chink left 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
formation of pollen; it comparatively rarely
(query ever) happens that the back or lower
surface of the antheral leaf is specially devoted
to the formation of pollen. On the other hand,
in cases like those of the common honuseleek,
where we meet with petaloid organs combining
the attributes of anthers and of carpels, we find
the inner layers devoted to the production of pollen,
the outer to the formation of ovules.

That the pollen-lobes are not to be taken as halves of a staminal leaf,
but rather as specialised portions of it, not necessarily occupying half
its surface, is shown also in the case of double-flowered Malvaceae, in
which the stamens are frequently partly petal-like, partly divided into

Fic. 157. Peta-
loid stamen of
Viola, with four
projecting plates.

PETALODY. 2993

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,' in which the connective

Fie. 158.—Portion of a double columbine (Aquilegia), 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

Fic. 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. Miiller’s paper on the anther of
Jatropha Pohliana, &c., referred to at page 255.

: See C. Morren, “On Spur-shaped Nectarines,” &e., ‘Ann. Nat.
Hist.,’ March, 1841, p. 1, tab. 11.

294, METAMORPHY.

anther-lobes are usually atrophied, and little or no
pollen is formed.

An occurrence of this nature in Tacsonia pinnati-
stipula, in conjunction with the partial detachment of
the stamens from the gynophore, led Karsten to estab-
lish a genus which he called Poggendorfia.*

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 (Polypetale,
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
spirallyarranged while the carpels are grouped in whorls,
the former will be more liable to petalody than the
latter, and vice versd. It has been before remarked,
that this condition is far more common in plants whose
petals, &c., have straight veins, lke 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.’ 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 Andrecium of Menizelia, &c., in

Seemann’s ‘ Journal of Botany,’ vol. iii, p. 209, and vol. iv (1866), p. 273
(Potentilla, &c.).

PPETALODY. 295

mens affords a satisfactory explanation of the appear-
ance in some double Malvacew, wherein the tufts of
adventitious petals are very liable to be mistaken for
buds, produced by axillary prolification in the axils of
the petals, but which are in reality compound and
petaloid stamens. At other times, however, true
axillary prolification exists in these flowers; but then
the supplemental florets have always a calyx, which is
wanting in the other instances.

Petalody of the stamens has been met with most
frequently in the following genera :

J *Ranunculus! /Z Asculus!

‘fori PRO

/ *Anemone!

/2*Geranium!

*Papaver !—2Q

,3* Pelargonium

Z= J fete

/ *Clematis! 3 *Tropxolum ! ee a
/ *Hepatica ! 7 /#Oxalis ! aI ih
/ *Ficaria! /3 *Impatiens! ys
/ Thalictrum. -,*Camellia! 7 {TA Ceee
J *Caltha! aay Thea! - Bit Ath AL
/ *Trollius ! /6Trifolium! 9 =
4 *Nigella! 7&6 Medicago ! yer 4. tA then
/ *Aquilegia ! /6* Ulex! bcc 3
/ *Delphinium ! / Spartianthus. 7 EvetLaiwen et =
/ *Adonis! / » Clitorea.
/ *Peonia! / Pisum! § Varlnetae.
3 *Nelumbium ! /6 Orobus ! A f
4 *Nymphea! / © Genista! 7. Cann pepplay i,
5S *Berberis! ‘ Spartium ! ao , v
*Paparert— /(-Cytisus! 46 Cena tenr
2. *Chelidonium! /GAnthyllis. +; Wtplrnecac
A Sanguinaria. /@Coronilla. A
S~ Podophyllum. /o Lotus! he Attra. om
é@ *Mathiola! ¢7*Rosa ! esa Ath
6 *Cheiranthus! 0) eKerria ! /3 Z y vy “e
& *Iberis! » *Spirea! / ¥ Ojai fatcas
G *Cardamine! » *Fragaria ! S35) Batloeanewne
t& *Hesperis , » *Potentilla! tb PEPSI
& *Barbarea ! » *Crategus ! P i
& *Sinapis! » Cydonia. 17)
*Brassica ! « *Pyrus! 46 Mapthrnnqeas
*Helianthemum ! » Eriobotrya! ;
¥* Viola! » *Amygdalus!
7 *Diantbus! » *Prunus!
4 *Saponaria ! 2@Myrtus!
¢ *Lychnis! 47*Punica! — Perce a
7*Silene ! ¥*Philadelphus !
/ OSagina ! / ¥*Deutzia!
/7* Hibiscus ! *Fuchsia !
+F Althea! Godetia !} 28 brary retire
//*Malva! Clarkia!
3F_
79
3
the

SIE Portulaca ! *Digitalis! > 4
Vn wash oriatts -Ribes! eo *Linaria !
PR OLE EGS Saxifraga! Z / Veronica !
Vy TIS. P- Daucus. 22 Calceolaria! ‘‘
Rip Ny heed sip, EXOree Achimenes. 367
eee nase, HOCTISSA | ' Gloxinia! 47
tte Mebieceta*. Gardenia!) _ Clerodendron !~O
Foun’ te / Lonicera! <© Bignonia. </
Srp Sambucus. 2 © Cyclamen! # 2
aif Viburnum. 22 Mirabilis. 4 2.
Scabiosa. Laurus! ¥ 3
*Campanula !2 ¢ Gladiolus! =<”
2S Platycodon ! * Crocus !
a8 BY ER. Calluna! Agszc Tris! a
ef Azalea ! ; *Galanthus! # 2
5 Rhododendron ! + Leucojum !
herrea hdtece *Arbutus! ' Sternbergia! «
2 *Hrica ! Hippeastrum. «
“ *Anagallis! 3 2 *Narcissus! os
Nemeo *Primula! 3 2 *Orchis ! a)
F ; * Jasminum ! = Catasetum ! // As
writs = Svringa! z= Hydrocharis. 4 7
a *Vinca ! Asphodelus. ¥ &
— *Nerium ! *Tulipa! "
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"ff *Calystegia! 24 Fritillaria!
» MaawtAre4eL“€, Convolvulus! ., *Tilium !
are Ipomea. + *Hyacinthus! “
bs Ahir *Datura! 3 & *Polianthes! # 9_
*Petunia! =. *Hemerocallis ! #9
a A at Solanum! 2... *Colchicum ! a
-— Orobanche. » “¥ *Sagittaria! #9
VAv 0 hae; Gentiana. 29° *Tradescantia !4 ©
‘A tia Mimulus. Commelyna! +,
. ; *Antirrhinum ! Tofieldia. ¥
CAN ; Gratiola !
Lb gHftt-yt Ct

296

METAMORPHY.

Petalody of the pistils—'T'aken 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, &e.,
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

PETALODY. 297

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-
loped.

In double flowers of Prinuwla sinensis it frequently
happens that the capsule is either partially leafy or
partly petal-hke; 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 followmg lst 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. Amyegdalus !

*Papaver ! Lonicera !

*Dianthus ! Scabiosa.
Saponaria ! Adschynanthus !
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 Crucifere.’

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,’ li, p. 351, tab. 12, Sinapis.

298 METAMORPHY.

point of view. OC. Morren has 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.’ A
similar change may be seen in the double Oleander.

Staminody of the bracts—An instance of this has been
already alluded to in Adies 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 Licopoh met with a similar
substitution of anthers for bracts in Melianthus major.’

Staminody of the sepals and petals——In the first named this
is of very rare occurrence. M. Gris has recorded an
instance in Philadelphus speciosus’ 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 ale and carina of the
corolla were thus changed.‘ There is in cultivation a
form of Savxifraga granulata wherein the petals are
replaced by stamens, so that there are fifteen stamens.
A similar change has been observed in Capsella bursa-
pastoris.

Cramer figures and describes a stamen occupying
the place of a petal in Daucus Carota.’ Turpin’ de-
scribes a similar occurrence in Monarda fistulosa, m
which the lower lip terminated in an anther, but this
may have been a case of adhesion. Moquin cites
from Chamisso, Digitalis purpurea, and from Jussieu,
Asphodelus ramosus, as haying presented this change,
and Wiegmann’ has seen anthers developed on the
awns of Avena chinensis. In semi-double flowers of
Ophrys aranifera and Orchis mascula, the lateral petals

1 «Bull. Acad. Belg.,’ xvii, part i, p. 516, ¢. 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.

1 «Mem. Legum.,’ p. 44.

* ‘ Bildungsabweich, ‘ Pflanz. Fam..,’ tab. 8, f. 12.

® « Atlas de Gothe,’ p. 55, t. 4, f. 18.
7 Wiegmann, ‘ Bot. Zeit., 1831, p. 5, tab, i.

STAMINODY. 299

are ocasionally partially antheroid, and others occur in
which two of the outer series of stamens, which are
ordinarily suppressed, are present, but in a petaloid
state. Reichenbach’ figures an illustration of this
change, and also Mogegridge.”

Staminody of the pistils—The existence of this change has
been denied by several authors, nevertheless, it is of
sufficiently common occurrence. Alexander Braun
notices the transformation of pistils into stamens in
Chives (Allium Scorodoprasum), and in which three
stamens appeared in the place of as many pistils, and
had extrorse anthers, while the six normal anthers are
introrse. In the horse-radish (Armoracia rusticana),
two of the carpels are frequently converted into stamens,
while two other organs absent from the normal flower
make their appearance as carpels. Roeper has observed
this phenomenon in Euphorbia palustris,’ and in Genti-
ana campestris. 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.’

Agardh has observed a similar thing in a hyacinth,
one half of the fruit of which contained seeds, and the
other half, anthers. B. Clarke mentions an instance
in Mathiola incana in which the carpels were disunited,
and antheriferous at the margin.®

The passage of pistils to stamens in willows has been
frequently remarked, as in Salix babylonica, silesiaca,
cinerea, Caprea and nigricans. One of the most curious
illustrations of this transformation in this genus is
given by Henry and Macquart (Erst. Jahrb. des bot.
Vereines am m. et n. Rhein., 1837). In the flowers in

1 ‘Je, Flor. Germ..,’ xiii, tab. 112, cecelxiv, f. 2.

Seemann’s ‘ Journal of Botany,’ 1867, p. 317, t. 72, A (Ophrys).
‘Enum. Euphorb.,’ p. 53.

‘ Linnea,’ i, p. 457.

‘De Balsam,’ p. 17.

2
3
4
6 B. Clarke, ‘ Arrangement of Phenog. Plants,’ p. 23.

300 METAMORPHY.

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 im the more perfect states were raised on three
filaments.

Campanula persicifolia, C. rapunculoides, and CO.
glomerata have been observed to present an anther
surmounting the pistil.t 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 imto anther-like
organs. Mohl has recorded an analogous malforma-
tion in Chamerops humilis, and in which the three
carpels were normally formed, and only differed from
natural ovaries in this, that along the two edges of
the ventral suture there was a yellow thickening,
which a cross section of the ovary showed to be an
anther-lobe filled with pollen.’

In Tofieldia calyculata a similar substitution of a
stamen for a carpel has been observed by Klotsch,'
and Weber* gives other instances in Prunus and Peonia.
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
Ricinus communis as having been in one instance

1 See ‘ Engelmann,’ p. 26, tab. 3, f. 10, 11, 14.
> « Ann. Sc. Nat.,’ ser. 2, t. viii, 1837, p. 58.
: ‘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
Peonia where the carpel was open and petaloid, and bore an anther on
one margin, and four ovules on the other. :

STAMINODY. 301

antheriferous.' Moggridge figures a flower of Ophiys
insectifera in which the rostellate process was replaced
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.

Staminody of the accessory organs of the flower—The scales
that are met with in some plants, either as excrescences
from the petals, or as imperfect representatives of
stamens or other organs, are occasionally staminoid ;
thus the scales of Saponaria officinalis, of Silene,
Neriwm Oleander, the rays of Passijflora, the corona of
Narcissus, have all been observed occasionally to bear
anthers.” 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. incompara-
bilis, and N. montanus.

M. Bureau found in some flowers of Antirrhinwm
majus two petal-like bodies standing up in front of, or
opposite to the two petals of the upper lip,‘ and similar
developments m 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 *Kuphorbiaceae,’ p. 205.

2 Seemann’s ‘ Journ. Bot.,’ iv, p. 168, tab. 47, f. 1.

3 Moquin-Tandon, 1. c., 220, Passiflora. Masters, ‘Journ. Linn. Soc.,’
1857, p. 159, Saponaria. Seemann’s ‘Journ. Botany,’ vol. ii, p. 107,
Narcissus.

+ * Bull. Soc. Bot. Fr.,’ 1857, p. 452.

302 METAMORPHY.

Pistillody of the perianth—The passage of the segments
of the perianth into carpels has been observed fre-
quently in Tulipa Gesneriana, the change in question
being generally attended by a partial virescence. M.
Gay is said by Moquin to have observed a flower of

Fria. 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 ‘Gardeners’ Chronicle,’ 1866, p. 897,

PISTILLODY. 303

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
cases the whole of the stamen appears to be changed,
while in others it is the filament alone that is altered,
the anther being deficient, or rudimentary; while, in a
third class of cases, the filament is unaffected, and
the anther undergoes the change in question. In
those instances in which the filament appears to be
the portion most implicated, it becomes dilated so as
to resemble a leaf-sheath rather than a leaf-stalk, as it
does usually.

One of the most curious cases of this kind is that
recorded in the ‘ Botanical Magazine,’ (tab. 5160, f.
4) as having occurred in Begonia
frigida already alluded to, and in
which, in the centre of a male
flower, were four free ovoid ovaries
alternating with as many stamens.
In the normal flowers of this plant,
as is well known, the male flowers
have several stamens, while in the
female flowers the ovary is strictly
inferior, so that, in the singular
flower just described, the perianth
was inferior instead of being 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 |
mae et of the oran ph ae ae

Se, numerary carpels in
called by the French “ bigarades ge a. poate
cornues,’’ the thalamus of the flower, pistils pa eine
which is usually short, and termi-
nated by a glandular ring-lke disc, is prolonged into
a little stalk or gynophore, bearing a ring of super-
numerary carpels. These carpels are isolated one

304 . METAMORPHY.

from another, and are formed by the transformation
of the filaments of the stamens.'

The additional carpels in the case of the apple of St.
Valéry, in which the petals are of a green colour, lke
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.)? 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 combinmg
the characters of both.’ Dunal and Campdera have
described flowers of [ames crispus, with seven pistils,
occupying the place of as many stamens.

In Papaver bracteatuwm a considerable number of the

Fig. 162.—Substitution of carpels for stamens in Papaver.

Stamens sometimes become developed into _pistils,
1 Maout, ‘ Lecons Element.,’ vol. ii, p. 488.
2 Poiteau and Turpin, ‘ Arb. Fruit,’ t. 37, and Trécul, ‘Bull. Soe. Bot.
France,’ vol. i, p. 307.
3 Clos, ‘Mem. Acad. Toulouse,’ 5 ser., vol. iii.

PISTILLODY. 305

especially those which are nearest to the centre of the
flower, and in these flowers the filaments are said to be-
come the ovaries, while the anthers are curled so as to
resemble stigmas. <A similar change is not infrequent
in Papaver somniferum. Goeppert, who found numerous
instances of the kind in a field near Breslau, says the
peculiarity was reproduced by seed for two years in
succession.’ Wigand (‘ Flora,’ 1856, p. 717) has noticed
among other changes the pistil of Gentiana Amarella
bearing two sessile anthers. Polemoniwm cerulewm is
another plant very subject to this change. Brongniart*
describes a flower of this species in which the stamens
were represented by a circle of carpels united to each
other so as to form a sheath around the central
ovary. By artificial fertilization M. Brongniart ob-
tained fertile seeds from the central normal ovary as
well as from the surrounding metamorphosed stamens.

Chevranthus Cheiri has long been known as one of
the plants most subject to this anomaly. De Candolle
even mentions it in his ‘ Prodromus’ as a distinct variety,
under the name of gynantherus. Brongniart (loc. cit.)
thus refers to the Cheiranthus :—‘‘ Sometimes these
six carpellary leaves are perfectly free, and in this case
they spread open, presenting two rows of ovules
along their inner edges, or these edges may be soldered
together, forming a kind of follicle like that of the
columbine; at other times, these staminal pistils are
fused into two lateral bundles of three in each bundle,
or into a single cylinder which encircles the true pistil.
In a third set of cases these outer carpels are only
four in number, two lateral and two antero-posterior,
all fused in such a manner as to form around the
normal pistil a prism-shaped sheath, with four sides
presenting four parietal placentz, corresponding to
the lines of junction of the staminal carpels.

1 « Bot. Zeit.,” 1850, t. viii, pp. 514, 664. ‘Flora,’ (B. Z.) 1832, t. xv,
p. 252; also cited in ‘Ann. des Serres et des aut dins,’ vi, pp. 241-5. See
also Schlechtendal, ‘ Bot. Zeit.,’ 1845, t. 3, p. 6

‘Bull. Soc. Bot. France,’ t. viii, p. 453.

20

306 : METAMORPHY.

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

Fie. 163.—Chetranthus Cheiri, var. gynantherus. a. Sepals and petals
removed to show carpellodic stamens. 0b. The same laid open. e.
Transverse section. d. Plan of flower with four carpel-like stamens, Xe.

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.’

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

1 See also Allmann, ‘ Rep. Brit. Assoc.,’ July, 1851.

PISTILLODY. 307

shows one of the intermediate organs from these
flowers, in which half the structure seems devoted to
the formation of ovules, while the other half bears a
one-celled anther. Lindley’ has also
described a case of this kind in a
species of Amaryllis.

In Savifraga 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, m the centre of
the flower. In this particular in-
stance there is usually no interme-
ciate condition between the stamen
and the pistil. Guillemin’ also de-
seribes a transformation of the stamens Fre. 164.—Struc-
into carpels in Huphorbia esula. ture half anther,

When the anther is involved it may *#!f ca"pel, ilvan.
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.’

In Rosa 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 Horticulture,’ ed. 2, p. 82.

2 «Mém. Soc. Hist. Nat. Paris,’ i, 16.

* Gen. Munro, ‘ Trans. Linn. Soc.,’ xxvii, p. 7.
* «Bull. Soe. Bot, Fr.,’ 1857, p. 21.

308 METAMORPHY.

Payer, in his ‘Organogénie,’ p. 38, mentions a
stamen of Dionewa bearing not only an anther, but
hkewise an ovyule.

Sempervivum tectorum and S. montanwin have long
been noticed as being very prone to present this change.
Mohl’ remarks that, in the transformation of the
stamens to the pistil in the common houseleek, the
filament of the stamen generally preserves its form,
the anthers alone undergoing change. At other times,
however, the transformation takes place at the same
time, both in the filament and in the anther. When
the stamens are numerous some of them remain in
their normal state, while others, and especially the
inner ones, undergo a change. Sometimes all the
stamens are changed simultaneously, while at other
times some of these organs may be found in which the
anther is partially filled with ovules, and partially with
pollen.

In the accompanying figures (fig. 165, a—h) a
series of intermediate stages is shown between the
ordinary stamen of Sempervivum tectorum and the
ordinary carpel, from which it will be seen that the
filament is little, if at all, affected, and that in those
cases where there is a combination of the attributes
of the stamen and of the pistil in the same organ the
pollen is formed in the upper or inner surface of the
leaf-organ, while the ovules arise from the opposite
surface from the free edge, (b, c, d, e, f, 9).

In a drawing made by the Rey. 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

' «Ann, Science. Nat.,’ t. viii, 1837, p. 50, and ‘Bot. Zeit.’ (R.), 1836,
t. xix, p. 513, &. See also MM. Sourd Dussiples and G. Bergeron,
‘Bull. Soc. Bot. France,’ viu, p. 349; Von Schmidel, ‘Icon. plant. et
Anal. part.,’ 1782, p. 210, fig. 54.

PISTILLODY. 309

into a long style, terminated by a flattened spathulate
stigma.

Delphiniwm elatum is one of the plants in which
this change has been most frequently noticed.’

In willows the change of pistils into staminal organs
has been frequently observed. In Salix babylonica
Prof. Schnizlem has described various transition stages

q h

Fic. 165.—Sempervivun tecotorum. a. Normal stamen. h. Normal
carpel. b,c, e,f,g. Structure partly staminal, partly carpellary. d.
Transverse section through c, showing pollen internally, ovules ex-
ternally.

between the carpels and the stamens, and in one
instance, in addition to this change, a perfect cup-
shaped perianth was present, as happens normally
in Populus.” Mr. Lowe also records the conversion
of stamens imto ovaries in Salir Andersoniana, and
this by every conceivable intermediate gradation.’

? Godron, ‘ Bull. Soc. Bot. Fr.,’ xiii, p. 82, Rev. Bibl.
* Cited in Henfrey, ‘ Bot. Gazette,’ in, p. 12.
5 «Ann. Nat Hist.,’ September, 1856, p. 56. See also Kirschleger,

310 METAMORPHY.

The following list will serve to show what plants
are most subject to this anomaly. It is difficult to
draw any accurate inference from this enumeration,
but attention may be called to the frequency of this
occurrence in certain plants, such as the Sempervivum,
the wallflower, the poppy, and the heath. Why these
plants should specially be subject to these changes
cannot be at present stated.

By the student of animal physiology such a change
as above described—equivalent to the substitution of
an ovary or a uterus for a testis—would be looked
on as next to impossible; the simpler and less spe-
cialised structure of plants renders such a change in
them far more easy of comprehension.

Thalictrum minus. Myrtus, sp.
Delphinium elatum. Campanula rapunculoides.
Magnolia fuscata. Polemonium ceruleum.
Bocconia cordata. Gentiana Amarella.,
*Papaver bracteatum ! *Hyrica Tetralix.
3 somniferum ! Stachys germanica.
nudicaule. Primula acaulis.
Dionza muscipula ! Rumex crispus.
Barbarea vulgaris. *Salix, sp. plur.!
*Cheiranthus Cheiri! Euphorbia esula.
Cochlearia Armoracia. Glochidion.
Tropeolum majus. Asphodelus ramosus.
Citrus Aurantium. Amaryllis.
*Sempervivum tectorum ! Lilium tigrinum!
montanum. longiflorum.
Begonia frigida! *Tulipa Gesneriana !
Cucumis, sp. var. cult. plurim.!
Cucurbita Pepo. Hemerocallis.
Pyrus Malus. Zea Mays.
Rosa arvensis ! Bambusa, sp.

Saxifraga crassifolia !

Pistillody 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. Zeit.),’ xxiv, 1841, p. 340, Salix alba. Henschel, ‘ Flora

(Bot. Zeit.),’ 1832, t. xv, p. 253, 8. cinerea. Hartmann, ‘ Flora (Bot. Zeit.),’
xxiv, p. 199, S. nigricans. Meyer, C. A., ‘ Bull. Phys. Math..,’ t. x, S. alba.

ce eel fl Bite (0
HETEROMORPHY.

THERE are certain malformations that have little in
common beyond this, that they cannot readily be
allocated in either of the great groups proposed by
writers on teratology. There are also deformities
which, unlike the majority of deviations from the
ordinary structure, are absolute and not relative.
While the latter are due to an exaggeration, or to an
imperfection of development, or, it may be, to a 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.

CHA P*RE-Re Ag
DEFORMITIES.

Tar 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-

De HETEROMORPHY.

trophy, Atrophy, &c., for malformations capable of
more rigid classification than those here alluded to.

Formation of tubes—T'he production of ascidia or
pitchers from the cohesion of the margms 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

! Fic. 166.—Portion of the under surface of a cabbage-leaf, with horn-
like excrescences projecting from it.

some portions as contrasted with others, whence arises
either a depressed cavity, as in the case of a leaf, or

ASCIDIA. 313

an expanded and excavated structure, when the stem
or some portion of it 1s 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-
bium, there is but little difference.

In cabbages and lettuces there not unfrequently
occurs a production of leaf-lke 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.

Fic. 167.—Lettuce leaf, bearing on the back a stalked cup, arising
from the dilatation of the stalk (?).

314 HETBROMORPHY.

In a variety of Codieum variegatum a similar forma-
tion may be seen to a minor extent. Hven the common
Scolopendrium vulgare occasionally produces small
pitchers of this character, as in the varieties named
perafero-cornutwm, Moore, and peraferum, Woll.’

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,’ in which
the leaves, epicalyx, sepals, and petals, were all provided
with tubular spurs.

In Cephalotus follicularis rudimentary or imperfect
pitchers may be frequently met with, im which the stalk
of the leaf is tubular and bears at its extremity a very
small rudimentary leaf-blade. It is not in all cases easy
to trace the origin and true nature of the ascidium, as
the venation is sometimes obscure. If there be a single
well-marked midrib the probability is that the case is
one of cohesion of the margins of the leaf; but if the
veins are all of about equal size, and radiate from a
common stalk, the pouch-like formation is probably
due to dilatation and hollowing of the petiole. Again,
when the result of a union of the margins of the
leaf, the pitcher is generally less regular than when
formed from the hollowed end of a leaf-stalk. Further
information is especially needed as to the mode of
development and formation of these tubular organs, so
as to ascertain clearly when they are the result of a
true cupping process, and when of cohesion of the
margins of one or more leaves. (See Cohesion, p. 31.
For “bibliographical references consult also A. Braun,
‘Flora v. Bot. Zeit.,’ 1835, t. xvii, p. 41, Aristolochia.)

Tubular formations in the flower—A similar formation of
tubes happens in some double flowers; for instance,
it is not infrequent in double flowers of Primula
sinensis, in which tubular petal-like structures are

' Moore, ‘ Nature Printed Ferns,’ 8vo edition, vol. ii, p. 154, et p. 173.
> «Flora (B. Z.),’ 1821, vol. iv, p. 717, ¢. tab.

FORMATION OF TUBES. Si

attached to the inner surface of the corolla; sometimes
these petaloid tubes replace the stamens, while at other

Fra. 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 haye 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,'
Tulipa Gesneriana, and occasionally on the sepals of

' Chavannes, ‘ Mon. Antirrh.’

316 HETEROMORPHY.

Fuchsia. They are very frequent in some seasons in
the corolla of certain calceolarias (C. floribunda). 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-

Fic. 169.—Corolla of tion seems to be purely irre-
gular tale sp pr: ae as ious] -
jecting from the lower lip. ’ 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, [Iinaria, &e. A similar formation of
conical out-growths may frequently be met with in the
fruits quite irrespectively of any disjunction of the
carpels.

Contortion—An irregular twisting or bending of the
stem or branches is by no means of uncommon 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
bemg 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

CONTORTION. 317

with an oblique form of the leaf. This state is some-
times present to an extreme degree, as in some varieties
of shrubs (Crataegus, Robinia, &e.) cultivated for their
singularly tortuous branches.

[J

Cc
)
2 df

Fria. 170.—Portion of the culm Fie. 171.—Portion of a branch of
of a Juncus, bent irregularly. Crategus oxyacantha, var. tortuosa.

Such cases as those just mentioned, however, are
but shehtly 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.

318 HETEROMORPHY.

M. Moquin-Tandon' alludes to a case of this kind in a
species of pine (Pinus), in which a branch ended in
four unequal divisions, which were strongly curved
from without inwards, then became united in pairs,
these latter in their turn blending into a single mass.

In the case of some beeches growing in the forest
of Verzy, near Rheims, the trunks of the trees are 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*® mentions the stems of Ituscus 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 versdé (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, Phanix dactylifera, origi-
nally observed by Goethe, and figured and described by
Jaeger ;*> 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 Thid., t. iv, 1857, p. 759.

3 Jaeger, “ De monstrosa folii Phanicis dactylifere conformatione a

Goetheo olim observata,” ‘ Act. Acad. Leop. Car. Nat. Cur.,’ vol. xvii,
suppl., p. 293, ¢. tab. color. iv.

SPIRAL TORSION. 319

as the leaflets increase in size, remaining from some
cause or other unbroken, and thus serving to restrain
the growth.

A similar irregularity of growth occurs, not 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.’ 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
Alstremeria, in Avena, and other plants. A similar
tendency is manifested in the flower-stalks, as in
Cyclamen and Vallisneria, and the whole inflorescence,
as in Spiranthes. Even the bark and wood of trees is
often disposed spirally. This is very noticeable in
some firs, and in the bark of the sweet chestnut
(Castanea), of Thuja occidentalis, and other trees. The
knaurs or excrescences which are sometimes found on
the roots or stems of trees afford other illustrations
of this universal tendency. These bodies consist of a
number of embryo buds, which, from some cause or
other, are incapable of lengthening. On examination
every rudimentary or undeveloped bud may be seen
to be surrounded by densely crowded fibres arranged
spirally.

The axes of nearly all twining plants are themselves
twisted, and twisted in a direction corresponding to
the spontaneous revolving movement exhibited by these
plants, as in the hop, the conyolvulus, passion flower,

' See Goethe, ‘ Ueber die spiral Tendenz.’

320 HETEROMORPHY.

&e., the degree of twisting being dependent to a great
extent on the roughness of the surface around which
the stem twines.’

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 line
round the stem, as in a specimen of Dracocephalum
speciosum described and figured by C. Morren. The
leaves of this plant are naturally rectiserial and 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 3 hence comes regularly into

3 *) Thus, the leaves in assuming
a new phyllotaxy, take one quite analogous to the
normal one.

One of the most curious instances that have fallen
under the writer’s own observation occurred in the
stem of Dipsacus fullonwm. (See ‘ Proceedings of

the : series a

1 See Darwin “On Climbing Plants,” ‘Journ. Linn. Soc. Botany,’
vol. ix, p. 5.

SPIRAL TORSION.

the Linnean Society,’ March 6, 1855,

BYAl

vol. 1, p. 370).

The stem was distended, and hollow, and twisted on

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 case,
disposed in a linear series, one over
the other, following the line of 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 poimt. 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-
quently in the species of Galium and
other Rubiacee.

Fie. 172.—Twist-
ed stem of Dipsacus

fullonum.

G. Franc’ was one of the first to notice this twisting
in Galiwm, and M. Duchartre,” 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 Galiwm Mollugo the branches

1 «Ephem. Nat. Cur.,’ dec. 2, ann. 1, 1683, p. 68, fig. 14.
2 * Ann. des Scienc. Nat.,’ third series, vol. i, 1844, p. 292.

21

a2? HETEROMORPHY.

are opposite in each verticil and crossed in the two
successive ones. ‘The stem is four-angled, each angle
haying a nerve. LHach 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 adjoming 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 Galiwm 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 amllary
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 1s 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 im the
way in which they were described by M. Duchartre,

SPIRAL TORSION. 323

but more commonly form a single continuous line;
when arranged in leaf-whorls it generally happens

Fic. 173.—Stem of Galiwm spirally twisted. From a specimen com-
muuicated 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 Hqui-
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 Reinsch’ cites
a case in Hquisetum 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

' « Flora,’ Feb. 4, 1858, p. 69, tab. ii, f. 3, and also ‘ Flora,’ 1860, p. 737,
tab. vii, f. 9.

324 HETEROMORPHY.

the two preceding, it was seen that the spire mceluded
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-
bound.

Fie. 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.

SPIRAL TORSION. 325

Morren spoke of this displacement of the floral organs
as ‘* speiranthie.’’?

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
Contortion.)

Hesperis matronalis. Mentha viridis.
Dianthus barbatus. Fraxinus vulgaris !
Pyrus Malus. Sambucus nigra.
torminalis, Zinnia.
Cercis siliquastrum! Phylica.
Punica Granatum. Beta.
Robinia pseudacacia ! Rumex, sp.
Rubia tinctorum. Ulmus campestris.
Dipsacus fullonum ! Casuarina rigida.
pilosus. Abies excelsa !
Gmelini. Lilium Martagon!
Scabiosa arvensis. candidum.
*Valeriana officinalis! *Asparagus Officinalis !
dioica! Sagittaria sagittifolia.
Galium aparine! Epipactis palustris.
* Mollugo ! Triticum repens !
verum ! Lolium perenne!
Hippuris vulgaris! Phleum pratense.
Veronica spicata. Juncus conglomeratus !
longifolia. Scirpus lacustris.
Hyssopus officinalis. Equisetum Telmateia.
Thymus Serpyllum. limosum.
Lamium purpureum ! fluviatile.
Dracocephalum speciosum. arvense !

Mentha aquatica.

Among the more important papers relating to this
subject may be mentioned :

Moquin-Tandon, ‘ El. Ter. Veg.,’ p. 181. Kros, ‘De Spira in plantis
conspicua.’ Morren, ‘ Bull. Acad. Roy. Belg.,’ 1851, tom. xviii, part i,
p. 27. Milde, ‘Nov. Act. Acad. Leop. Carol. Nat. Cur., 1839. Ibid.,
vol. xxvi, part ii, p. 429, Equisetum. Irmisch, ‘ Flora,’ 1858, t. i,
Equisetum. Vrolik,‘ Nouv. Mem. Instit. Amsterdam,’ Liliwm. 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, ¢. tab.

326 HETEROMORPHY.

Veget., t. i, p. 155, tab. xxxvi, Mentha, &c. Alph. de Candolle, ‘ Neue
Denkschr. Allg. Schweiz. Gesellschft.,’ band v, tab. vi, Valeriana. Du-
chartre, ‘Ann. Sc. Nat.,’ ser. 3, vol. i, p. 292. ‘ Gardeners’ Chronicle,’
July 5, 1856, p. 452, ¢. ic. wylogy., spiral branches from Guatemala—
tree not known.

Spiral twisting of the leaf 1s scarcely of so common oc-
currence as the corresponding condition in the stem.
In Alstremeria it occurs normally, as also in some
erasses. In the variety annularis of Salix babylonica
the leaf is constantly coiled round spirally. A similar
contortion occurs in a variety of Codicewm variegatum
lately introduced from the islands of the South Seas
by Mr. J. G. Veitch.

Fern fronds are occasionally found twisted m the
same manner, e.g. Scolopendriwm vulgare var. spirale.

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 Hodgsonia, Strophanthus, &e.

M. Decaisne’ found a flower of the melon in which
one of the segments of the calyx was prolonged into
a tendril, and Kirschlegér 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 ito
a tendril.

In Cobea 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 Hecremocarpus
scaber. On the other hand, in the vine, the axial nature
of the tendril is revealed by the not infrequent presence
of flowers or berries on them, as also in Modeeca and
some Passifloracee.

1 Moore, ‘ Nature-printed Ferns,’ 8vo edition, vol. u, 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, ii, p, 546.

INTERRUPTION. 327

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
cement.

Interrupted growth—This term is here used in the
same sense as in ordinary descriptive botany, as when
an ‘‘interruptedly pinnate’ leaf is spoken of. A similar
alternation may be observed occasionally as a terato-
logical occurrence, though it is not easy to account
for it.

Fig. 175 shows an instance of the kind in a radish,

Fic. 175.—Interrupted growth Fie. 176.—Interrupted growth
of Radish (from the ‘ American in Apple.
Agriculturist.’)
and fig. 176 a similar deformity in the case of an
apple, the dilatation of the flower-stalk below the
ordinary fruit producing an appearance as if there were
two fruits one above another.

In leaves this peculiar irregularity of development
is more common.

328 HETEROMORPHY.

In some varieties of Codiewm variegatum the leaves
resemble those of Nepenthes, as the basal portion is
broad, and terminates in a projecting midrib destitute
of cellular covering, and this again terminates in a
small pouch or pitcher. Somewhat similar varia-
tions may be found in ferns, especially Scolopendrium
vulgare.

Instead of the pouch there is formed sometimes in
the plant last mentioned a supplementary four-lobed
lamina, the four lobes being in two different planes,
and diverging from the midrib, so that the section
would resemble }<, the poimt 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. Miiller,
p. 295.

Cornute leaves (Molia 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 Codiewin varieqatum already
referred to.

Flattening.—'T'here are some plants whose stem or
branches, instead of assuming the ordinary cylindrical
form, are compressed or flattened; such are some species
of Hpiphyllum, Coccoloba, Bauhinia, &c. The same
thing occurs in the leaf-like branches of Ruscus, the
flower-stalks of Xylophylla, Phyllanthus, Pterisanthes.
Martins proposes to apply the word ‘cladodium’ to such
expansions, just as the term phyllodium is applied to
the similar dilatation of the leaf-stalks. If we exclude

POLYMORPHY. 329

instances of fasciation, 7. e. where several branches are
fused together and flattened, we must admit that this
flattening does not occur very often as a teratological
appearance.

Mr. Rennie figures and describes a root of a tree
which had become greatly flattened in its passage
between the stones at the bottom of a stream, and had
become, as it were, moulded to the stones with which
it came into contact.’

The spadix of Arwm, 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,
&e.,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.’

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 C. linearifoliwn,
in which, owing to the suppression of the lamina, the
petiole had become dilated into a ribbon-like expan-
sion—déformation rubanée of Moquin.

CHAPTER II.
POLYMORPHY.
Usvatiy 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

' Loudon’s ‘Magazine Nat. Hist.,’ vol. ii, p. 463.
* C. Morren, ‘ Bull. Acad. Belg..,’ "1852, t. xix, part ili, p. 444.

330 HETEROMORPHY.

dimensions as another; one flower resembles very closely
another flower of the sameage andsoon. 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 apples. The variation in form is usually

HETEROPHYLLY. 331

dependent on a greater or less degree of lobing of the
margin of the leaf; thus,in the yellow jasmine, almost
every intermediate stage may be traced from an ovate
entire leaf to one very deeply and irregularly stalked.
Broussonettia papyrifera, and Laurus Sassafras, and the
species of Panaz, may be mentioned as presenting

Fic. 177.—Syringa persica laciniata, showing polymorphous leaves.

this condition. Sometimes in the last-named genus,
as also in Pteridophyllum, every gradation between
simple and compound leaves may be traced. The
horse-radish (Cochlearia Armoracia) may also be in-
stanced as a common illustration of polymorphism in

Bee HETEROMORPHY.

the leaves. In ferns it is likewise of frequent occur-
rence, markedly so in Scolopendrium D Urvillei, in
which plant every gradation from a simple oblong
frond to an exceedingly divided one may be found
springing from the same rhizome at the same time.

A similar protean state, but little less remarkable,
occurs in many of our British ferns, notably m Scolo-
pendriwin vulgare, of which Mr. Moore enumerates no
fewer than 155 varieties, 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,
&e. Most of these variations are mentioned under
the head of the particular morphological change of
which they are illustrations.

The effect of a change in the conditions of growth
in producing diversity in the form of the leaf may
be here alluded to. Ficus stipulata, a plant used to
cover the walls of plant-stoves in this country, and
growing naturally on walls in India, like ivy, produces
leaves of very different form, size, and texture, when
grown as a standard, from what it does when adhering
to a wall. Marcgraavia wmbellata furnishes another
example of a similar nature, as indeed, to a less extent,
does the common 1

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 s 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.

DIMORPHISM. 333

Pteris quadriaurita, in which the fronds emerging from
an adventitious bud are very different from the ordinary
fronds.

—S-

x S\ SE IS Ma i. z R
iF) oS ; % = » ~

fe

PAT re

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—T'his 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’ (Solanwm tuberosum), which produces first double
and sterile flowers, and subsequently single fertile ones ;
the other portions of the plant do not differ much.

1 * Gard. Chron.,’ 1845, p. 790.

334. HETEROMORPHY.

Stackhousia juncea, according to Clarke, has mixed
with its perfect flowers a number of apetalous blossoms
destitute of anthers.’

This peculiarity is well exemplified in the tribe
Gaudichaudiee of the order Malpighiacee. <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. Violacee, Campanulacee, &e.
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
Veronica.’

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 Phenog. Plants,’ p. 36.
2 «Bull. Soe. Bot. France,’ 1856, t. i, p. 569.

DIMORPHISM. oD

are really sexual forms of one and the same species,
ordinarily occurring on different plants, 7. ¢. dicecious,
but occasionally formed on the same spike. The same
excellent observer has demonstrated that the di- and
trimorphic forms of Primula, of Linwm, Lythrwm, and
other plants—forms differing maimly in the relative
length of the stamens and styles, are also connected
with striking differences in the number of perfect seeds
produced. The most perfect degree of fertility is
obtained when the stigma of one form is fertilised by
the pollen taken from stamens of a corresponding
height. On the other hand, when the union is, as Mr.
Darwin states, illegitimate, that is, when the pollen is
taken from stamens not corresponding in length to the
style, more or less complete sterility ensues in the
progeny, sometimes even utter infertility, ‘such as
happens when two distinct species are crossed, so
that, in point of fact, the offspring of these illegitimate
unions correspond almost precisely to hybrids.’

Mere variations of form arising from hybridisation
or other causes hardly fall within the lmits 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’ 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 his work
on the ‘ Variation of Animals and Plants,’ vol. ii, p. 181.
2 Loc. cit., 1, 336.

336 HETEROMORPHY.

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.—'l'hese curious departures from
the normal form can only be mentioned mceidentally in
this place, as they pertain more to variation than to
malformation.

The occasional production of shoots bearing leaves,
flowers, or fruits of a different character from those
found on the normal plant, is a fact of which gardeners
have largely availed themselves in the cultivation of
new varieties. The productions in question have been
attributed to various causes, such as cross-breeding,
grafting, budding, dissociation of hybrid characters, or
reversion to some ancestral form, all of which 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 Cytisws Adan, the
trifacial orange, &c., are discussed.

wy
wo
NI

ALTERATIONS OF COLOUR.

CHAPTER III.

ALTERATIONS OF COLOUR. !

CHANGES in the colour of the several organs of plants
are more often either pathological or the result of
variation than of malformation properly so called.

Alterations in colour arise from a diminished or an
increased amount of colouring matter, or from an 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
Richardia 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

organs.
22

338 ALTERATIONS OF COLOUR.

Virescence—Engelmann’ 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’ 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; mstead
of falling off speedily they become persistent when so
affected.

Some flowers are more lable to virescence than others.
The common honeysuckle, Lonicera Periclymenum, is
one of these, and it is noticeable in this plant that the
calyx remains unaffected—a circumstance which Morren
says shows the distinctness of virescence from 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-
hike 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

1 ‘De Antholys,’ p. 32, § 38.
2 «Bull. Acad. Belg..’ xvii, part 2, p. 151, c. tab.

CHROMATISM. 309

abortive anthers, and the pistil also is in a rudimentary
condition. Umbellifere are not unfrequently subject to
this change, e. g.,Vorilis Anthriscus, Daucus Cavota, Hera-
clewm Sphondylium, Carum cari, &e. Pr imaulaceee, again,
are frequently subject to virescence. Among Composite
the followmeg species are recorded as having had green
flowers—Cirsiwm tricephalodes, Senecio vulgaris, Calen-
dula officinalis, Pyrethrum Parthenium, Carduus erispus,
Hypocheris vadicata, Hieracium prealtwm, Cirsium
arvense, Coreopsis Drummondi.' In Ranunculacee vires-
cence has been observed in Delphinium elatwin, 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 coronaria 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,
‘Verhandl. des Bot. Vereins. Brandenb.,’ heft 1, 2, Anchusa. Christ,
‘ Flora,’ 1867. pp. 376, tab. 5, 6, Stachys.

340 ALTERATIONS OF COLOUR.

the petals, the others bemg of their ordmary green
colour.

The occurrence of coloured bracts, as in Poinsettia,
Bougainvillea, &c., is very common under natural con-
ditions, and need not here be further alluded to.

Increased intensity of colour often accompanies
teratological changes ; an instance has just been alluded
to in the Gesnera: the feather hyacinth, Muscari
comosum, furnishes another illustration, the adven-
titious pedicels being brightly coloured.

In fasciated stems, also, of herbaceous plants, 1t 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, 1s 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.’

EOCy Bone Pt.
DEVIATIONS FROM THE ORDINARY NUMBER OF ORGANS.

To a certain extent the number of the organs of a
plant is of even greater consequence for purposes of
classification than either their form or their arrange-
ment; 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, m the minor subdivisions, and
especially where the absolute number of parts is large,
considerable variation may occur, so that descriptive
botanists frequently make use of the term indefinite,
and apply it to cases where the number of parts is
large and variable, or, at any rate, not easy to be
estimated.

Considered teratologically, the changes, as regards
the number of organs, are readily grouped into those

342 NUMBER.

consequent on a decreased and into those resulting
from an increased development. The alteration may
be absolute or relative. There may be an actual
deficiency in the number of parts or an increase in
their number, but in either case the change may be
simply a restoration of the primitive number, a species
of peloria, in fact. An increased number of parts,
moreover, may depend not so much on the formation
of additional parts as on the subdivision of one.

It seems also desirable to treat separately those cases
in which there is an increased number of buds either
leaf-buds or flower-buds, as the case. may be, as hap-
pens in what is termed prolification. This formation of
buds occurring, as it does, often in unwonted situations
is treated of tnder 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),

Ip Aoki Ee,
INCREASED NUMBER OF ORGANS.

AN augmentation in the number of parts may arise
from several causes, and may sometimes be more
apparent than real. True multiplication exists simply
as a result of over-development ; the affected organs
are repeated sometimes over and over again each in
their proper relative position, and without any 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, owimg 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, followimg 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 splittimg of one petal, for
instance, into several ;—somewhat in the same manner
as one may separate successive layers of tale one from
the other. :

344 INCREASED NUMBER.

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 Primulacee,
wherein a second row of stamens is occasionally present
in the adult condition, and renders the floral symmetry
perfect.

The double daffodil, where there are from forty to
fifty petaloid organs instead of fifteen, and wherein
each piece exhibits a more or less perfect coronal lobe
at the junction of the claw and the limb, has been cited
as an objection to chorisis, though it is difficult to see
on what grounds.

In Delphiniwm, as shown by Braun,’ 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
referrible to an absolute increase in the number of
whorls, and not to chorisis.’

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 sphtting of an organ
already perfectly formed mto two or more portions,
either in the same plane as the original organs,

? Braun, ‘Pringsheim Jahrbuch f. Wiss. Bot.,’ 1858, 1, p. 307, tab. 22, 23.
° Henfrey, ‘Jour. Linn. Soc. Bot.,’ vol. ii, p. 159

CHORISIS. 345

** parallel chorisis ;”’ or at right angles to it ‘‘ collateral
chorisis.”’ Indeed, before so much attention had been
paid to the way in which the floral organs are deve-
loped, it was thought that an actual splitting and
dilamination did really take place ; Dunal and Moquin
both assert as much. The truth would rather seem to
be that, im the so-called parallel chorisis at least, the
process is one of hypertrophy and ovyer-deyelopment
rather than of splittmg. 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. Out orowths from leaves, multiplymg
the laminar surface, are alluded to under the head of
hypertrophy, and it is probable that some of the cases
of duplication of the flower, or of the formation of
adventitious segments outside the ordinary corolla as
alluded to in succeeding paragraphs (see Pleiotaxy of
the corolla), are due to a similar process.’

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 httle or no doubt that there
are in reality six stamens in Orchidacee, of which one
only, under ordinary circumstances, is developed.
When the numerical symmetry is restored, as it some-
times is, it 1s obvious that the augmentation that
_occurs is of a different character from that arising from

1 On the subject of chorisis or dédoublement the reader may profit-
ably consult Moguin- Tandon, ‘Ess. sur les Dédoublements,’ and the
same author in ‘ Ann. Sc. Nat.,’ t. XXVil, p. 236, and ‘ El. ies Veget.,’
p. 337. Dunal, ‘Consid. Org. Fleur.,’ Montpell., 1829, p. 32, note 3.
A. de St. Hilaire i in ‘ Ann. Se. Nat.,’ ser. 3, t. i1, p. 355, ae Lindley,
‘Elements of Botany,’ p. 76. Asa Gray. ‘Botancal Text Book.’

346 INCREASED NUMBER.

a repetition or renewed development of organs. When
the increased number arises from multiplication proper,
or from repetition, the ordinary laws of alternation are
not interfered with, but if from chorisis or ‘*‘ dédouble-
ment,” it may happen that the normal arrangement 1s
disturbed.

Without studying the mode of development, it is
not in all cases possible to tell under which of the
above categories any particular instance should be
placed ; hence, in the following sections, except where
otherwise stated, the cases are grouped according to
the appearance presented in the adult condition, rather
than to the way in which the changes from the typical
condition are brought about. With reference to the
foliar organs it is necessary to distinguish those cases
in which there is, from any cause, an augmentation in
the number of component parts of a whorl, from those
in which the increase takes place in the numbers of the
whorls themselves.

CHAPTER I.

MULTIPLICATION OF AXILE ORGANS, INFLORESCENCE, ETC.

By Linné 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, cew instar nidi
Pice, quod vulgo a genio ortum arbitratur; frequens
apud nos in Betula, presertim Norlandie, in Carpino
Scanie, nec infrequens in Pinu.’

By some of the older authors this condition was
called polyclady. In some cases, it would seem to

1 «Phil Bot.,’ § 274.

AXILE ORGANS. 347

be due to fungi as in the witches’ brooms (hexenbesen)
of the German forests; in other instances, itis a result
of mutilation as after the operation of pollarding.
Moquin-Tandon' 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
*onaurs”’ or “ burrs,” met with in elms, &c., also in
certain varieties of apples, are clusters of adventitious
buds, some of which might, and sometimes do, lengthen
out into branches.
An increased number of branches also necessarily
arises when the flower-buds are replaced by leaf-buds.
Occasionally, a great increase in the number of
pedicels, or flower-stalks, may be met with in conjunc-

Fic. 179.— Flower stalks of Bellevalia comosa, nat. size, after Morren.

1 «El. Ter. Veget.,” p. 392.

348 INCREASED NUMBER.

tion with a decreased number of flowers, as in the
wig-plant (Ithus Cotinws), or the feather-hyacinth
(Bellevalia comosa). In these cases the supernumerary

Fre. 180.—Tuft of branches at the end of the inflorescence of Belle-
valia comosa, enlarged after Morren.

pedicels are often brightly coloured. 'T'o this condition
Morren gave the name mischomany, from pisyoc, a
pedicel, a term which has not generally been adopted.’

M. Fournier’ describes a case in the butcher’s broom
(Ruscus aculeatus), wherein from the axil of the mmute
leaf subtending the flower a secondary flattened
branch proceeded.

Duchartre® cites the case of a hyacinth which, im
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 stems which become adherent to the primary
flower-stalk, cannot, in all cases, be determined. Cer-

1 «Bull. Acad. Belg.,’ xvii, part ii, p. 38.

2 * Bull. Soe. Bot. Fr.,’ vol. iv, 1857, p. 760..
3 [bid., vol. viii, 1861, p. 159.

AXILE ORGANS. 349

tainly, in some cases examined by me the latter was
the case.’

Under this head, too, may be included those cases
wherein an ordinarily spicate inflorescence becomes
paniculate owing to the branching of the axis and the
formation of an unwonted number of secondary buds.

Fie. 181.—Increased number of male catkins in the hazel Corylus
avellana.

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. J aeger figures and describes a
buneh of Pinus sylvestris bearing i 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.”

M. Reichardt describes an analogous case m the
‘same species, and attributes the ordinate number of
cones to a fungus (Peridermium pint). In this ease

1 a Gard. Chron.,’ July, 1866, p. 656, and Clusius, ‘ Plant. Rar.,’
lib. 2, p. 143, Tulipa serotina wodvedadyc, minor, &e. ‘ Hort. Eysttett.
Plant. Ver.,’ fol. 12.

2

* ‘Jaeger de Pini sylvestris monstrositate,’ Stuttgardt, 1828.

300 INCREASED NUMBER.
there were no less than 227 cones, but each one half
the size of the ordinary cones.’

Of asimilar 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 pecuharity, as in the Hgyptian
wheat or corn of abundance (Triticum compositum),
certain varieties of Maize, etc. Similar exuberant
erowths occur in Orchidacee, in Cyperacee, e.g. Carea,
in Restiacew, 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 Equisetacee a similar plurality of spikes
occurs often as a result of mutilation.” 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 Leucoiwm
estivum described by M. Gay.® Irmisch described a
similar phenomenon in LZ. vernwm; and Mr. Moggridge
has communicated drawings of a similar formation im
the same species grown in the neighbourhood of
Mentone.

1 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, ‘Knollen und
Zwiebelgew.,’ tab. 7, figs. 10, 11.

AXILE ORGANS. Som

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.—Brocecoli, 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
erasses 1s also met with under some circumstances. I
have seen this in Hordewm and Lolium, and an instance
is figured in Avena by Dr. Wiegmann.’ M. Duval Jouve’
records a similar occurrence in Catabrosa aquatica, the
spikelets of which contained from two to seven flowers.’

1 «Flora, 1831, p. 5, tab. 1; see also Hanstein, ‘ Flora,’ 1857, p. 513.
Schlechtendal, ‘ Bot. Zeit.,’ xviii, p. 381.

2 «Bull. Soc. Bot. Fr.,’ ix, p. 8.

3 Tt 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

302 MULTIPLIGATION.

CHAPTER IL.

MULTIPLICATION OF FOLIAR ORGANS.

Tur 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 ee 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 1s 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 verticillata. In
one instance the pseudo leaf divided, and from the division proceeded a
little axis, bearing at its summit a verticil of pseudo leaves. This 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.

POLIAR ORGANS. Soo

surface of which project, at right angles to the primary
plane, other secondary leafy plates; but these are,
strictly speaking, cases of hypertrophy (see Hyper-
trophy).

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 1s greater than usual.

The increased number of leaves in a whorl may well
be designated as “‘ polyphylly,” using the word in the
same sense as in ordinary descriptive botany, while
*‘pleiotaxy”? may be applied to those cases in which
the number of whorls is increased.

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 leafletsin 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

\ A \\ \A

Fic. 183.—Supernumerary leaflet, Ulmus campestris.

produced in the elm. In the one case the new leaflet
springs from the apex of the petiole and partially fills
23

B04 MULTIPLICATION.

the space consequent on the obliquity of the base of
the leaf. In the other it would seem as if two distinet
leaves emerged from the stem in juxtaposition. This

NS se
Qn AW
SRA

SS
WS

Ss

Fie. 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 im
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

FOLIAR ORGANS. ove

tubular, horn-like projections, and in the centre a
thread-like process representing the midrib and termi-

Fie. 185.—Supernumerary leaf of hazel.

nated by a small two-lipped limb.’ Dr. Ferdinand
Miiller speaks of a leaf of Pomadervris elliptica as bearing
a secondary leaf on its under surface.’

The leaves of Heterocentron macrodon have likewise
been observed occasionally to produce leaflets from
their upper surface.

To this production of leaves from leaves the late
Professor Morren applied the term ‘ autophyllogeny.’”
The Belgian botanist figures a small perfect leaf sprie-
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

' «Bull. Soc. Bot. Fr.,’ vol. vii, 1860, p. 587.
> «Fragment. Phyt. Austral.,’ part xx, p. 270.
* «Bull. Acad. Belg.,’ xvi, pt. i, p. 60, “ Fuchsia,” p. 125, ¢. ic.

356 MULTIPLICATION.

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.’

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 (Trifoliwm 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. Linné, who im 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 7. repens
bore seven leaflets. Schlechtendal alludes to a similar
increase in number in Cytisus Laburnwm, 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. Taberneemontanus
‘Krauterbuch,’ §, 222. Schlechtendal, ‘ Bot. Zeit.,’ ix, p, 083, xiv, p. a.

1 “Théorie de la feuille,”’ ‘ Av ch. des Aono Bibl. Tae rs 1868,

FOLIAR ORGANS. S57

Maugin, ‘Bull. Soc. Bot. Fr.,’ 1866, t. xiii, p. 279. See also Cramer,
‘ Biidungsabweich,’ p. 92. Walpers, ‘Linnea,’ 1840, p. 362 (7-leaved).
Schlechtendal, ‘ Bot. Zeit.,’ 1844, p. 457, Cytisus. Wigand, ‘ Flora,’ 1856,
p- 706,

Frondiferous leaves have much the appearance of
branches provided with leaves, and they may be com-
pared with those instances in which an adventitious
bud is placed on the surface or edges of the leaves, as
in Gesnera, Cardamine, &c. In truth, the two conditions
merge one into the other, as in some begonias, where
the ramenta often become leaf-like and bear small
bulbils in the axil.

When frondiferous leaves die the appendages die
also, but when a true bud has been formed on a leaf it
does not of necessity die with the leaf that bears it, but
separates from it and continues to grow independently.

Increased number of stipules, spathes, &e—Seringe relates
the occasional presence of two or three additional
stipules upon the leaf-stalks of Salix fragilis, and even
makes a variety (Salix pendula, var. multistipulata).

An increase in the number of the spathes has been
often noticedin Arads.' Prof. Alex. Braun has studied
this subject in some detail.” In Calla palustris the
shoot which continues the growth of the plant proceeds
from the axil of the last leaf but one; the very last leaf
producing no bud, but if accidentally a shoot is developed
in this latter situation it produces flowers at once. No
leaves are formed, but, on the contrary, two or three
spathes surround the spadix, so that the presence ofan
increased number of spathes in this plant is associated
with the development of a side shoot from the axl
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

' See Engelmann, ‘De Antholysi,’ p. 16, section 12.
* «Verhandl. des Botanisch. Vereins Brandenburg,’ 1859, 1 heft.

358 MULTIPLICATION.

occur in other Arads as Arum maculatum, Richardia
ethiopica, and Anthurium Scherzerianwm, frequently
combined with a leaf-like appearance of the spathes
and sometimes with a subdivision of the spadix imto
two or three branches.

Engelmann relates the occurrence of an increased
number of glumes in Bromus velutinus associated with
suppression of the flowers.

Polyphylly—As previously explained, this term is
here applied to those cases in which the members of
any particular whorl are increased in number, the
whorls themselves not necessarily bemg 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—T'his 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, Xe.

Among plants with normally opposite leaves the
following occasionally produce them in whorls of
three :— Lonicera brachypoda, L. Xylostewm, Weigela

rosea, Cornus mas, Vinca nunor, &e.

Paris quadr ifolia may frequently be met with with

five leaves in its whorl, or even six.'

Increased number of bracts——This is not of infrequent
occurrence; one of the most curious instances is that
recorded by Mr. Edwards? in Cerastiwm glomeratum,
where, in place of the usual pair of bracts at the base
of the head of flowers, there was a whorl of six or eight,
forming an involucre. The flowers in this case were
apetalous and imperfect.

Polyphylly of the calyx——This may occur without any

- 1 See Henslow, ‘Mag. Nat. Hist.’ 1832, vol. v, p. 429.
7 « Phytologist,’ Se enka 1857.

POLYPHYLLY. 359

other perceptible change, while at other times the
number of the other parts of the flower is proportion-
ately increased. In a flower of a plum six sepals in
place of five sometimes exist; a precisely similar
occurrence in the flowers of the elder (Sambucus), the
Fuchsia, and of Gnanthe crocata, may occasionally be
met with. In the latter case, indeed, there are some-
times as many as ten segments to the calyx, and
this without the other parts of the flower being
correspondingly augmented. Among monocotyledons
a similar increase is not uncommon, as in Tulipa,
Allium, Iris, Narcissus, &c.

In some plants there seems to exist normally much
variation in the number of parts; thus in some species
of Lacistema im adjacent flowers the calyx may be
found with four, five, or six segments.

Most of these cases of polyphylly affecting the calyx
may be explained by lateral chorisis or fission.

Polyphylly of the corolla—This may happen in 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 suppressed ;
thus, when in aconites we meet with four or five horn-
like nectaries (petals) instead of two only, as usual, the
supernumerary ones are accounted for by the inordinate
development of parts which ordinarily are in an abortive
or rudimentary state only. This is borne out by what
happens in Balsaminee. Inthe common garden balsam
the fifth petal is occasionally present, while in Hydrocera
trijlora this petal is always present.

In a flower of a Cyclamen recently examined there
were ten petals in one series, the additional five bemge
evidently due to the subdivision of the five primary
ones; the natural circular plan of the flower was here
replaced by an elliptical one. A similar occurrence
takes place in the flowers of maples (Acer), which
sometimes show an increased number of parts in their

360 POLYPHYLLY.

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 ordimary con-
formation.’

I am indebted to Mr. Hemsley for the communication
of a similar specimen in O. apifera, in which there
were two divergent lips, each with the same peculiar
markings. One of the sepals in this flower was
adherent to one of the lateral petals. This 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
_ (see p. 380).

The following enumeration will suffice to show the
genera in which an increased number of petals or
perianth-seoments in any given whorl most frequently
occurs.

Anemone! Solanum.
Ranunculus! Veronica.
Aconitum ! Cyclamen !
Raphanus. Primula !
Bunias. Anagallis !
Saponaria. Plumbago.
Dianthus ! Jasminum.
Pelargonium ! Syringa !
Hibiscus. Tradescantia.
Fuchsia. Tris.
Sarothamnus ! Tigridia.
Lotus! Narcissus.
Ulex! Tulipa.
Prunus! Convallaria !
Trifolium. Paris !
Cinanthe and Umbellif. pl. ! Hyacinthus !
Sambnueus ! Allium !
Bryonia. Ornithegalum.
Campanula. Orchidew, sp. pl. !

—E———

1 Seemann’s ‘ Journal of Botany,’ iv, p. 168, t. 47, f. 3.

pn

STAMENS. 361

For other illustrations see multiplication of whorls, petalody ; see also
Mogquin, loc. cit., p. 350. Engelmann, loc. cit., p. 20, § 18. Cramer, loc.
cit., p. 25.

Polyphylly of the andrecium—An increased number of
stamens frequently accompanies the corresponding
alterations in other whorls, and seems, if anything, to
be more frequent among monocotyledonous plants than
among dicotyledonous ones; thus, we occasionally find
tetramerous flowers in Crocus, Hyacinthus, Tulipa, Iris,
Tigridia, &c., and more rarely in Yucca (Y. flevilis').

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 peioria as
in Iinaria, 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 Lamiwm, Mentha, Chelone;? 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, Hichler’ figures a flower
of Matthiola 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* 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’ 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* mentions the presence

1 * [lust. Hortic.,’ 1866, misc., p. 97.

2 See Fresenius, ‘Mus. Senkenb.,’ bd. 2, p. 43. Schlechtendal, ‘ Bot.
Zeit.,’ iv, pp. 403, 492, Veronica tetrandra.

3 « Flora,’ 1865, tab. 6, fig. 8.

* «Org. Veget.,’ t. i, p. 497, pl. 42, f. 3.

° «Bl. Ter. Veg.,’ p. 354.

° Cited in “ Rev. Bibl.” of ‘Bull. Soc. Bot. Fr.,’ 1866, p. 171.

362 POLYPHYLLY.

of seven stamens in another Crucifer, /onopsidiwm
acaule. Here the supernumerary organ was placed
between two of the long stamens. The effect of
chorisis in producing an augmentation of parts 1s well
seen in some plants that have some of their flowers
provided with staminodes or abortive stamens, and
others with clusters or phalanges of perfect stamens.
Thus, in the female flowers of Liquidambar there are
five small staminodes without anthers, whereas in the
male flower the stamens are numerous and grouped
together in phalanges, so that the relation of simple to
compound stamens is in this case readily seen, as also
in many Malvacew, Sterculiacew, Buttneriacee, Tilacee,
and Myrtacee. 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, m
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 eles by Moquin from Dunal’ goes
far to support the notion here adopted. The majority
of the stamens of laurels (Laurus) have, says M. Dunal,
on each side of the base of their filaments a small
glandular bifid appendage ; these excrescences are lable
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
Sstamen, so that in these latter flowers there are twelve
stamens.

1 Loc. cit., 351.

GYN®CIUM, 363

M. Clos' mentions a flower of rue (Jéuta) wherein
there were two stamens joined together below and
placed in front of a petal, as in Peganwm.

Buchenaw* mentions a flower of Lotus uliginosus in
which there were eleven stamens, namely, two free
and nine monadelphous; and Hildebrand describes an
analogous increase in a flower of Sarothamnus scoparius
in which, in conjunction with a seven-toothed calyx,
there were two carinas and fourteen stamens. It
would seem probable in this case that there was a
coalescence of two flowers at an early date and conse-
quent suppression of some of the parts of the flower.
Whether this was the case or not in this particular
Ulustration, 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 gynecium.—Moquin® 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 Antirrhinum, where there are usually only two carpels

1 *Mém. 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. 304.

364 POLYPHYLLY.

present, but where, under peculiar circumstances, five
may be found—thus rendering the symmetry complete.’
In Papilionacew, wherein usually only one carpel is
developed, we occasionally find two, or even more, as
in Wistaria, Gleditschia, Trifolium, &e. In Prunus and
Amygdalus from two to five carpels are occasionally to
be found,’ in Mimosa five, in Umbellifercee three to five ;
in some composites, e.g. Spilanthes, five carpels have
also been noticed; in Crucifere three and four, in
orasses three.” 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 Crucifere, Umbelli-
fere, and Inliacew, are the orders most frequently
affected. Cases of peloria are not included in the
subjoined list.

Nigella. Ptelea.
Aquilegia. Citrus !
Peonia ! Philadelphus.
Delphinium ! Prunus!
Tberis. Amygdalus !
Diplotaxis. Crategus !
Lunaria. Fuchsia !
Ricotiana. Trapa !
Octadenia. Cassia.
Draba! Cercis.
Lepidium. Medicago.

*Cheiranthus ! *Phaseolus !
Dianthus. Wistaria.
Brassica ! Gleditschia.
Parnassia. Affonsea.

*A cer ! Trifolium !

! Giraud, ‘ Hd. Phil. Mag.,’ Dec., 1839.
* See Cerasus Caproniana, D. C. ‘ Plant. Rar. Hort. Genev.,’ tab. 18.
3 Nees, ‘ Linnea,’ v, p. 679, tab. 11 (Scheenodorus).

POLYPHYLLY. 365

Archidendron. Coleus.
Mimosa. Veronica !
Robinia. *Digitalis !
Diphaca. Antirrhinum !
Ccesalpinia. Linaria.
Vicia. Gloxinia !
Anthyllis. Symphytum.
Cucurbita. Anchusa.
Passiflora ! Polygonum.
Sambucus! Euphorbia.
*(@nanthe! Cneorum.
Daucus ! Mercurialis !
Angelica ! Chenopodium.
Heracleum ! Sueeda.
Silaus. Beta.
Carum. Corylus !
Thysselinum. Lambertia.
Campanula! Cocos!
Spilanthes. Tigridia.
Chrysanthemum. Tulipa!
Anagallis. Tris!
Primula ! Narcissus !
Fraxinus ! Album !
Lycium. Ornithogalum.
Cobea. Gagea!
Datura! Tradescantia !
Solanum ! Schcenodon.
Sesamum. Bambusez.
Sideritis.

A few additional references may here be given to papers where an
increased number of carpels i is described :—Hngelmann, ‘ De Antholys,’
§ 17, p.19. Bernhardi, ‘ Flora,’ 1838, p. 129. Schkuhr., ‘ Bot. Handb.,’
t. 179. Godron, ‘Ann. Se. Nat.,’ ser. 5, vol. 1, p. 280, tab. xviii,
pluricarpellary Crucifers. Weber, ‘ Verhandl. Nat. Hist. Vereins. Rhein.
Pruss.,’ &c., 1860, Cerasus, &e., &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, Bambusee. Alph. de Candolle, ‘Neue Denkschrift, ;
Cheiranthus. Schimper, ‘Flora,’ 1829, 1, p. 433. Wigand, ‘Bot. Unter-
such.’ Fleischer, ‘ Missbild. Cultur Pfl’ Cramer, ‘ Bildungsabweich,’
p. 65, Umbellifere.

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 1t must be remembered that, in general,
the augmentation is not confined to one whorl, but
affects several; thus, if the sepals are increased, the

366 MULTIPLICATION.

petals are likely to be so likewise, and so forth. One
of the most curious illustrations of this is that recorded
by Mr. Berkeley’ 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.

Fic. 186.—Plum. Increased number of parts in the calycine, corol-
line, and carpellary whorls respectively.

In Primulacee this general augmentation has been
frequently noticed.”

Among Orchidee the instance related by Dr. Seu-
bert is worth alluding to here. This botanist observed
and figured a flower of Orchis palustris with tetra-
merous arrangement of parts, that is to say there were

1 «Gard. Chron.,’ 1852, p. 452. ie
* See Cramer, ‘ Bildungsabweich,’ pp. 16, 24, *

OVULES. 367

four outer segments to the perianth, four petals, of
which two were lip-lke, four stamens, three of which
were rudimentary, and an ovary with four parietal
placentze.’

The following list will serve to show in what plants
this general augmentation of parts has been observed
most frequently :

Ranunculus. Sambucus !
Clematis ! *Primula !
- Delphinium. Anagallis !
Brassica ! Lycium.
Ruta. Solanum.
Acer ! Symphytum.
Prunus ! Syringa !
Rosa ! Linaria.
Rubus. Chenopodium.
Philadelphus ! *Paris !
Chrysosplenium Convallaria !
Umbellifere, sp. pl. ! Allium.
*Fuchsia ! *Tnilium !
-Ginothera. *Tulipa !
Adoxa. Ornithogalum.
Bryonia. *Gagea !
Cucumis ! Tradescantia !
Campanula! Orchidezx, sp. pl. !

Increased number of ovules or seeds——'I'his appears not to
be of very frequent occurrence, at least in those plants
where the number of these organs is normally small;
where, as in Primula, the ovules and seeds are 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-

* * Linnea,’ 1842, p. 389, e. ie.

368 MULTIPLICATION.

ceous condition of the carpel, in which the margins
are disunited. In such cases the ovules may occupy
the margin or may be placeda short distance within it,
as in the case of some open carpels of Ranuneulus
Ficaria,' 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 Tyi-
folium vepens becomes foliaceous (see Frondescence),
the outer ovules are generally two or more instead of
being solitary. So, also, in the Rose with polliniferous
ovules (see p. 274). Among Umbellifere 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 oyule is itself imperfect.”

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. 582, 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 contimued
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 be added Daucus, Heraclewm, Ke.

EMBRYOS. 369

nary circumstances cease to grow (see ante, p. 364). In
the 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
paragraph.

Increased number of embryos—A ripe seed usually con-
tains but a single embryo, although in the ovular state
preparation 1s commonly made for more; and, indeed,
mn certain natural orders. plurality of embryos in the
same seed does occur, as in Cycadew and Conifere. In
the seeds of the orange (Citrus), in those of some Huphor-
biacee, &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.!

Plurality of embryos has also been observed in—

Raphanus sativus. *Viscum album !
*Citrus Aurantium ! Daucus Carota.
Diosma, sp. Ardisia serrulata !
Hypericum perforatum. Cynanchum nigrum.
Triphasia aurantiaca. fuscatum.
* Asculys Hippocastanum ! Euphorbia rosea.
Euonymus latifolius. Ceelebogyne ilicifolia.
*Mangifera indica! Allium fragrans.
Eugenia Jambos. Funckia, sp.
Amygdalus vulgaris! Carex maritima.
Vicia, 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.
24,

370 MULTIPLICATION.

See Schaner’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, lke
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 acwminata 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 J examined bore out the latter view, as mm
the same pod were seeds with two, three, and four coty-
ledons respectively.

Ihave also seen three cotyledons present in embryo-
plants of Correa, Crataegus Oxyacantha, Dianthus sinensis,
Daucus Carota, Cerasus Lauro-cerasus. De Candolle
alludes to a case of the kind in the bean, and figures
a species of Solanwm with three cotyledons.’ Jaeger
alludes to a similar instance in Apiwm Petroselinwm ;”
Ehrenberg to one in the marigold (Calendula) ;* 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.* This fusion seems frequently
to accompany increase in the number of cotyledons.
It was so in the Correa and in the Crategus 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,’ in some

1 §Organog. Veget.,’ tab. 53. 2 * Missbild.,’ p. 206.

’ Ehrenberg, ‘ Flora,’ 1846, p. 704. 4 * Flora,’ 1860, tab, 7.
5 “Ann. Sc. Nat.,’ 3 ser., t. x, p. 207. :

PLELOTAXY. 371

Coniferze, which he considers to be improperly termed
polycotyledonous. Whether this holds good in the
Loranths, where (Nuytsia, Psittacanthus) an appearance
of polycotyledony exists, is not stated. In the case of
the rue (Ruta) figured by M. A. de Jussieu’ 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
polyphylly and pleiotaxy, as here proposed.

Pleiotaxy in the bracts—An increase in the number of
bracts has been met with very constantly in a species
of Mesa, and in a peculiar variety of carnation, called
the wheat-ear carnation.” 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 Amarella, where the bracts are increased in

1 ¢Mem. Mus.,’ xii, t. 17.

2 ‘Nov. Act. Acad. Nat. Cur.,’ xv, tab, xxviii, f.3; ‘ Bot. Mag.,’ t. 1622.
“Caryophyllus spicam frumenti referens.” A similar malformation in
Dianthus barbatus is not uncommon. It has lately been introduced into
gardens under the name of Dianthus “mousseuzx,” but is not likely to
find favour with gardeners.

ove PLEIOTAXY

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.

ff
MI
nA
Nr MEW
i 9 Me | y i

Fig. 187.—Wheat-ear carnation. The appearance is due to the
multiplication of the bracts and the suppression of the other parts of
the flower.

It has been noticed also in the common pea, Piswm
sativum, and M. Lortet' records a case of the kind
in rica nultiflora, 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.

OF THE BRACTS. 373

usual, and were covered for their whole length with
small rose-coloured bracts arranged in irregular whorls,
the upper ones sometimes enclosing imperfect flowers.
In the ‘ Gardeners’ Chronicle,’ 1865, p. 769, is figured
a corresponding instance of Delphiniwm Consolida, in

Fra. 188.— Delphiniwm Consolida. Fia. 189.—Multiplication of
Multiplication of bracts at the ex- bracts, &c., Pelargonium.
pense of the other parts of the
flower.

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 Pelargoniwm 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
malformation.

374 PLEIOTAXY.

Cornus mas and C. suecica sometimes show a triple
involucre.! Irmish? 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’®
mentions as having produced bracts in unwonted
numbers, Lythrwm Salicari ia, Plantago major, Veronica
spicata, Hehiwm vulgare, Melilotus arvensis, and Rubus
fruticosus.

It must here be remarked that this great number of
the bracts occurs naturally in such plants as Godoya,
in which the bracts, or, as some consider them, the
seoments 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 Roeper observed a calyx con-
sisting of a double series, each of five sepals, in con-
junction with other changes. It is also common in
double columbines, delphiniums, nigellas, &e.

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

1 Weber, ‘ Verhandl. Nat. Hist. Vereins. Rhein. Pruss.,’ 1860.
2 * Bot. Zeit.,’ 1848, p. 217.

3 *De Anthol.,’ p. 17, § 12.

4 «Tinnea,’ vol. ii, 1827, p. 85. ®

PERIANTH, 375
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 tothe 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. Valéry 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’ cites the following plants as occasionally
presenting a repetition of the calyx, in most cases with
a suppression of the other floral whorls :—Stachys lanata,
Myosotis palustris, Veronica media, Aquilegia vulgaris,
Nigella damascena, Campanula rapunculoides.

Pleiotaxy in the perianth—Increase in the number of
whorls in the perianth is common in lilies, narcissus,
hyacinths, &c. It may be also met with occasionally
among orchids. The lily of the valley (Convallaria
maialis) seems also to be particularly subject to an
increase in the number of parts of which its perianth
consists, the augmentation being due partly to repe-
tition or pleiotaxy, partly to the substitution of petaloid
segments for stamens and pistils.”

In this place may also be mentioned the curious
deviation from the ordinary structure occasionally met
with in Liliwm candidum, and known in English gardens
as the double white lly. In this case there are no
true flowers, but a large number of petal-like segments

' «De Antholysi,’ p. 17, tab. ii, 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 Antholysi,’ p. 18,
&e. For similar changes in Gagea arvensis see Wirtgen, ‘ Flora,’ 1838,
t. xxi, p. 350, and ‘ Flora,’ 1846, p. 353. Some of these are cases of
synanthy.

376 PLEIOTAXY OF THE

disposed in an irregular spiral manner at the extremity
of the stem, some of the uppermost being occasionally
verticillate.’

Fria. 190.—Double white lily. Multiplication of perianth-segments
and other changes.

Pleiotaxy of the corolla—With reference to double
flowers, it was remarked by Linné that polypetalous
flowers were, as he said, multiplied, while monopetalous
flowers were duplicated, or triplicated, as the case may
be,” a statement that is true in the main, though it

! Schlechtendal, ‘ Bot. Zeit.,’ xx, 1862, p. 301.
2 « Phil. Bot.,’ § 126. ’

COROLLA. S77

requires modification. In the case of polypetalous, or

rather dialypetalous flowers, the petals may be very
largely increased by multiplication, as im roses, ane-
mones, pinks, &c. In the last-named genus the number
is often so much increased that the ‘ealyx 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, 1t 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 Leguminose, ‘such as Ulex ewropeus 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."

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 rotundifolia,
Gardenia sp., Nerium Oleander, Serissa sp., Arbutus
Unedo, &e. The change is associated with petalody of
the stamens and pistils.

A more frequent change among the monopetalous
orders 1s 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.”

' C. Morren, ‘ Bull. Acad. Belg.,’ xix, part ii, p. 17.

3/78 PLEIOTAXY.

The same condition occurs frequently in some
species of Datwra and Campanula.

Fie. 191.—Campanula rotundifolia. ‘“ Double flowers” resulting from
dialysis and multiplication of the petals.

In Antirrhinwn imajus 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 Hrica hyemalis.

Mr. W. B. Hemsley has kindly furnished me with
flowers of a similar kind occurring in wild specimens of

ANDR@CIUM. 379

Epacris impressa,' and there are analogous phenomena
in the common honeysuckle (Lonicera Periclymenum),
in which three corollas and no stamens often occur.

This duplication may either be accounted for on the
theory of chorisis above alluded to, or by supposing
that the extra corolline whorl is due to a series of con-
fluent petalodic stamens; that the latter is the true
explanation, in certain cases at least, is shown by some
flowers of Datura fastuosa, m which the second corolla
was partially staminal in its appearance, and bore
nearly perfect anthers, in addition to the five ordinary
stamens, which were unaltered either in form or position.
Some partially virescent honeysuckle flowers have a
similar structure.

There are other cases of apparent multiplication or
duplication, due, probably, rather to the formation of
outgrowths from the petals than to actual augmentation
of their number. These excrescences occur sometimes
on the inner surface of the petals, or of the corolla;
at other times on the outer surface, as in some gloxinias,
&e. This matter will be more fully treated of under
the head of hypertrophy and enation.

Pleiotaxy of the andrecium—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 frequent where these organs are
arranged, not in verticils, but in one continuous spiral
line.

In Crucifere 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 Caryo-
plyllee, three and four verticils of stamens have been
met with. In Lonicera Periclymenwm a second whorl
of stamens more or less petalodic sometimes occurs.

Moquin mentions a variety of Rubus fruticosus in

‘ *Seemann’s Journal of Botany,’ iil, p. 354,

380 ANDR@CIUM.

which nearly 900 petaloid organs existed in the place
of the twenty-five or thirty stamens natural to the
plant, the other organs of the flower being in their
ordinary condition, with the exception of the pistil,
which did not attain its full size. Baillon records the
occasional existence of two rows of stamens in Ditaxis
lancifolia.

Increased number of stamens in orchids, &e— 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 androecium in this family
are so important in reference to what is considered its
natural conformation, that it seems desirable, in this
place, to enter upon the teratological appearances 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

ORCHIDACE’. 381

irregular, consisting not only of a petal, but of two
abortive stamens incor porated 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 Cypripedium, while the ordinary sta-
men of the outer row is deficient, two of the mner
series are present. The diagram, fig. 192, will serve

Fie. 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 2 should have been 1. (See text.)

to show the arrangement of the parts as above de-
scribed. + 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; a1, a 2,

382 ANDR@CIUM OF

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. 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
observer.

Mr. J. T. Moggridge has described and figured a flower of Ophrys
insectifera in which there was a vestige of a second stamen present,
probably one of the inner series fig. 192 (a ?).2 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.*

Reper is stated by Cramer’ to have seen a specimen of Orchis morio
with two stamens.

In a flower of Habenaria chlorantha, described by the late Professor
Henslow,° 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. ui, p. 183a, and in Darwin, ‘ Fertilisation of
Orchids,’ p. 292. See also Criiger, ‘ Journ. Linn. Soce.,’ t. viii, p. 134.

2 «Seemann’s Journal of Botany,’ vol. iv, p. 168, tah. 47.

* Thid., t. iv, 1866, p. 168, t. xlvii, f. 1. ‘

* « Bildungsabweich,’ p. 8; see also ‘ Bot. Zeit..’ 1852. p. 425.

* * Journ. Linn. Soc.,’ t. ii, p. 104, tab. 1, fig. B.

ORCHIDACE.A. 383

A flower of Cattleya violacea afforded a similar illustration; but in
this case only one of the inner stamens was developed, and this in the
form of a small petal, partly adherent to the column.

In Dendrobium normale, Falconer, not only is the perianth regular,
but the column is triandrous,' the three stamens (according to the
diagram of its structure given by Lindley) pertaining to the outer row.

In a specimen of Dendrobium hemoglosswm 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 a1, 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.” I have myself met with three such
flowers in the same species. The stamens present were A 1, w 1, a 2.

Dr. J. E. Gray exhibited at the Botanical Society of London, in
August, 1843, a specimen of Ophrys apifera with a triandrous column,
the supernumerary anthers belonging, apparently, to the inner whorl.

In his ‘ Catalogue of the Plants of South Kent,’ p. 56, tab. iv, f. 16,
the Rev. G. E. Smith describes and figures a flower of O. aranifera with
a triandrous column, seemingly of the same kind as that spoken of by
Dr. Gray.

Mr. Moggridge met with a triandrous flower in the same species, and
refers the appearance to “a fusion of two flowers, accompanied by
suppression and modification.”? 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* figures a flower of Mavillavia 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 (natwraliter

1 Lindl., “ Orchid. Ind.,” ‘ Jour. Linn. Soc.,’ iii, p. 9.

2 « Arch. Bot.,’ ii, p. 300, tab. xvi, f. 11.

° *Seemann’s Journal of Botany,’ v, p. 318, tab. Ixxii, figs. A 4, 4a.
4 “ Monstr. Veg.,” in ‘ Neue Denkschrift,’ p. 17, tab. vii.

384 ANDR@CIUM OF

conformate) in Orchis morio.! Richard, as cited by Moquin-Tandon,
Lindley, and others, describes and figures a peloria of Orchis latifolia
with regular triandrous flowers.”

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? 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 1t would appear as if the additional labella in this instance were
the representatives of two stamens of the outer whorl. Beer likewise
has put on record the existence of a triandrous Cattleya.*

A specimen of Catasetwin eburneum forwarded by Mr. Wilson Saunders
was normal so far as the sepals and two lateral petals were concerned,
but the anterior petal or labellum was flat and in form quite like the two
lateral ones; the column was normal and in the situation of the two
anterior stamens of the outer series A 2, A 3, were two labella of the
usual form (fig. 156, p. 291). Perhaps the Oncidiwm 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
(A2, A3). Within these was another perfect stamen occupying the
position of the anterior stamen of the inner series (43). In another
flower of the same species, gathered at the same time (fig. 194), there were
three sepals not at all different from those of the normal flower. The
three petals next in succession were also, in form and position, in their
ordinary state. In colour, however, the two upper lateral petals differed
from what is customary, in having the same purplish-brown tint which
characterises the lip. Within these petals, at the upper part of the
flower, there was the ordinary column, and at the opposite side, 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 (A2, A3). It is, perhaps, worthy of notice that the ar-

! Flora,’ t. viii, 1825, p. 736.

* «Mem. Soc. d’Hist. Nat.,’ ii, 1, p. 212, tab. iii.

3 * Bull. Soc. Bot. Fr.,’ t. vii, 1860, p. 26.

4 «Beitr. Morphol. und Biol. Orchid.,’ quoted by Cramer; ‘ Bildungs-
abweich,’ p. 9. :

ORCHIDACE®. 385

rangement of the coloured spots on the true labellum, and that on the
adventitious lips, replacing the two lower of the outer stamens, were

Fie. 193.—Diagram showing the arrangement of parts in a mal-
formed flower of Ophirys aranifera (see p. 384).

not of a similar character. The supernumerary lips had the z-shaped
marking which is so common in this species, while the true lip was,
as to its spots, much more like O. apifera. Alternating with this last
whorl were three columns, all apparently perfectly formed and differing
only from the ordinary one in their smaller size and corresponding to

Fic. 194.—Malformed flower of Ophrys aranifera with two super-
numerary lips and three additional stamens.

al,a2,a3. 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
25

380 ANDR@CIUM OF

positive sign of fusion in the bracts or other part of the flower, seemed,
however, to negative the idea of fusion.!

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 ophioglossoides 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 aranifera just mentioned. “The
peculiarities of these flowers,” writes Professor Gray, “are that they
have three labella, and that the column is resolved into small petaloid
organs. The blossom is normal as to the proper perianth, except that
the labellum is unusually papillose, bearded almost to the base. The
points of interest are, first, that the two accessory labella are just in
the position of the two suppressed stamens of the outer series, viz. of
A2 and a3, as represented in the diagram, fig. 192; and there is a small
petaloid body on the other side of the flower, answering to the other
stamen, Al. Secondly, in one of the blossoms, and less distinctly in
another, two lateral stamens of the inner series (a1 and a2) 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 Jsochilus is normally
triandrous.

A tetrandrous flower of Cypripedium has also been recorded.

In Isochilus, according to Cruger, there are often five stamens, and

Fra. 195.-—Diagram of flower of Orchis mascula with two additional
lips. two perfect and two imperfect stamens (after Cramer).

' Masters, ‘ Journ. Linn. Soc.,’ viii, p. 207. See also Rodigas, ‘ Bull.
Soc. Bot. Belg.,’ iv, p. 266, for similar changes in Cypripedium Hookere.

ORCHIDACE.AL. 387

there are several, besides those already mentioned, in which six more
or less perfect stamens have been seen—of these the following may
be taken as illustrations. A hexandrous flower of Orehis militaris has
been recorded by Kirschleger,’ and in the accompanying diagram (fig.
195), from Cramer,? 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.

Morren* describes some flowers of Orchis morio in which there were
three sepals, three petals, and within the latter two other ternary series
of petals; this would seem to be a case of petalody of all six stamens.
Morren, however, seems to have considered the additional segments as
repetitions of the corolline whorl, though he describes a central mass as
the column bearing a “ souvenir of the anther.” Nevertheless, there is
no decisive evidence either in his figure or his description in support of
his opinion as to the nature of the central mass, which might be a
distorted condition of the styles, or, as is more probable, a rudimentary
and irregular flower. Morren also describes another flower of the same
plant in which there were three sepals, two lateral petals partially lip-
like in aspect, a third labellum normal, two additional labella 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 homo-
logue. Thus it was in a specimen of Odontoglossum Alexandre 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, &c. Again, the existence
of adventitious stamens does not necessarily imply the development of
organs usually suppressed, inasmuch as they may result from the
assumption by the lateral petals of staminal characteristics.

Nevertheless, as far as teratology is concerned, specimens may be
found in which some or all of the usually suppressed stamens of Orchi-
dacee 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.
2 + Bildungsabweich,’ p. 11, tab. xiv, f. 3.
3 «Bull. Acad. Roy. Belg.,’ t..xix, part 2, p. 171.

388 PLEIOTAXY OF THE

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 gynecium—An increase in the number
of whorls of which the pistil consists 1s not of very fre-
quent occurrence. Generally after the formation of the
whorl of carpels, the energy of the growing point ceases,
or if by chance it be continued, the result is more 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 Delphiniwm, where the ovaries are arranged

Fig. 196.— Increased number of Fie. 197 —Fruxt of St. Valery
carpels, tulip. apple cut lengthwise.

GYN@CIUM. 389

in spiral series, is not strictly referable to the present
category.

The orange is one of the plants most frequently
subject to an augmentation in the number of carpellary
whorls; sometimes this is due to the stamens assuming
the guise of carpels, but at other times the increase
occurs without any alteration in the stamens or other
organs. If the adventitious carpels be exposed, they are
covered with yellow rind, while those portions that are
covered by the primary carpels are destitute of rind.
Some varieties of the double tulip are very subject to
a similar change, but, in this case, the petals and the
stamens very frequently become more or less 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 haying been
removed.

In the St. Valery apple, already referred to, there is
a second whorl of carpels above the first, a fact which
has been made use of to explain the similar structure
of the pomegranate.

The tomato (Lycopersicuin esculentum) is another
plant in which an adventitious series is frequently
produced, and generally in combination with the pri-
mary series.

In the Chinese primrose (Prinwla sinensis) 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’ mentions a flower of Gentiana pur-
purea with four carpels in one series, and five others
in the circle immediately above them. Wigand’ alludes
to an instance wherein there was a second pair of
carpels above the first in Vinca herbacea. Dr. Sankey
has forwarded flowers of a Pelargonium haying a double
series of carpels, eight in the outer row, five in the
inner, and this condition is stated to exist in the flowers

1 * Organogr. Végét., t. i, p. 509, tab. 40, figs. 6, 7.
> « Flora,’ 1856, p. 715.

390 PLEIOTAXY.

of the same plant for two years consecutively. In
Aquilegia T have met with a similar increase in the
whorls of carpels.’ Meissner records a similar aug-
mentation in Polygonwm orientale.’

Wigand’ 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. Wichler* has put on record a similar case in a
capparid.

Marchand’ mentions a polycarpellary berberid (Hpi-
medium 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 from a transformation, or substitution of
carpels for stamens.

Lastly, the instance cited by Dr. Allman® in Saai-
fraga Geum ray 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
forced. : :

In addition to the references already cited the follow-
ing may be given:

Duchartre, ‘ Ann. Se. Nat.,’ 4 ser., vii, p. 23 (Tulip).
Ferrari, ‘ Hesperides,’ pp. 271, 395, 405. Duchartre, ‘Ann. Se. Nat.,’

‘Linn. Trans.,’ t. xxiii, p. 3
‘Monog. Polygon,’ pl. 3, K

1 64, tab. 34, fig. 5.
2 f.

3 ¢ Plora,’ 1856, tab. viii.

4

Sole YN
Thid., 1865, tab. ix, f. 6.

> * Adansonia,’ vol. iv, 1864, p. 127. ¥

6 * Ann. Nat. Hist.,’ 1845, vol. xvi, p. 126.

PLELOTAXY. 391

4 ser., 1844, vol. i, p. 294. Maout, ‘Legons Elément.,’ vol. ii, pp. 488-9.
Clos, ‘ Ann. Se. Nat.,’ 1865, p. 317 (Citrus Aurantium).

Clos, ‘ Bull. Soc. Bot. Fr.,’ vol. xiii; ‘Rev. Bibl.’ p. 75. Pasquale,
‘Reddicont Accad. Sc. Fis. e Math. Napoli.’ Octr. 1866 (Solanum Lyco-
persicum).

On the general subject of multiplication, in addition
to previous citations, the reader may be referred to
A. P. de Candolle, ‘Théorie Elément. 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. 351). Thus spikelets of
wheat occasionally produce more than the three florets
which are proper to them.’ 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.

1 See Schlechtendal, ‘Bot. Zeit., t. xvii, p. 381 (Triticum); also
‘ Flora,’ t. xiv, 1831, p. 5 (Avena).

PAM eh Te

DIMINISHED NUMBER OF ORGANS.

A DIMINUTION in the number of parts is generally due
to suppression, using that word as the equivalent of
non-deyelopment. It corresponds thus in meaning
with the Fehlschlagen of the Germans, the avortement
complete of Moquin and other French writers. It
differs from atrophy, or partial abortion, inasmuch as
the latter terms apply to instances wherein there has
been a partial development, and in which evolution has
gone on to a certain extent, but has, from some cause
or other, been checked. These cases will be found under
the head of diminished size of organs. As the word
abortion is used by different authors in different ways,
_ it is the more necessary to be as precise as possible in
the application of the term. In the present work
abortion is used to apply to cases wherein parts have
been formed, but wherein growth has been arrested at a _
certain stage, and which, therefore, have either remained
in statu quo, while the surrounding parts have increased,
or have, from pressure or other causes, actually 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

SUPPRESSION. 393

obliterated by the pressure exercised by the constantly
increasing bulk of adjacent parts, or possibly has
become incorporated with them. In the adult flower
the appearances are the same, though the causes may
have been different.

CHAPTER I.
SUPPRESSION OF AXILE ORGANS.

ABSOLUTE suppression of the main axis is tantamount
to the non-existence of the plant, so that the terms
“acaulescent,” ‘‘acaulosia,’ &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, &e.

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

394 SUPPRESSION.

evidence furnished by teratology is conflicting, but
there seems httle 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 1s
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 1s 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-hke
projection from an axis arrested in its ulterior develop-
ment. ‘The secondary projections from the original one
correspond to an equat 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 1s
complete, and from which all possible primary projec-
tions are developed.’

If, as M. de Candolle considers, the leaf and the
branch differ merely in the fact that the vascular system
is complete in the latter, and partly atrophied in the
former, it would surely be better to consider the ‘‘ calyx-
tube”’ of the rose as a concave axis rather than as a
leaf, seeing that he admits the fibro-vascular system to
be complete in the case of the rose.

1 «Théorie de la feuille,’ p, 24.

a

FOLIAR ORGANS. 395

With reference to this point the reader is referred to
Mr. Bentham’s account of the morphology and homo-
logies of the Myrtacew in the ‘ Journal of the Linnean
Society,’ vol. x, p. 105. See also ante, pp. 71, 77.

Some doubts also exist as to the nature of the beak
or columella of such fruits as those of Geraniacer,
Malvacee, Umbellifere, Euphorbiacee, &e. 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
because, now and then, it happens that the organ in
question is completely wanting, and hence affords an
illustration of suppression.

CHAPTER II.

SUPPRESSION OF FOLIAR ORGANS.

THis subject may be considered, according as the
separate leaves of the stem or of the fiower are affected,
and according as either the number of members of
distinct whorls, or that of the whorls themselves, is
diminished.

The terms aphylly, meiophylly, and meiotaxy may
be employed, according as the individual leayes are
altogether wanting, or with reference to the diminished
number of parts in a whorl, or a decrease in the
verticils. .

Aphylly—Entire suppression of the leayes is a rare
phenomenon. Under ordinary circumstances it occurs
in most Cactacew, in some of the succulent Euphorbias,
and other similar plants, where the epidermal layers of
the stem fulfil the functions of leaves. But even in
these plants leaf-like organs are present in some stage
or another of the plant’s life.

396 MEIOPHYLLY OF THE

Partial suppression of the leaf occurs sometimes in
compound leaves, some or other of the leaflets of which
are occasionally suppressed. Sometimes, as Moquin
remarks, it is the terminal leaflet which is wanting,
when the appearance is that of Cliffortia, at other
times the lateral leaflets are deficient, as in Citrus or
Phyllarthron. Ononis monophylla and Lragaria 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.’

In lke manner varieties of the following plants
occur with simple leaves, ftosa berberifolia (Lowea),
Rubus Ideus, Robinia pseudacacia, Fraxinus excelsior,
Sambucus nigra, Juglans wigra, &e.

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
Stellate, and frequently in Paris quadrifolia, 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
' * Rev. Hortic.,’ 1866, p. 467.

COROLLA. 397

of sepals is not a very common occurrence among
dicotyledonous plants. Sermge figures a proliferous
flower of Arabis alpina with two sepals only, and a
similar occurrence has been noticed in Diplotaxis tenwi-
folia.

In Cattleya violacea the writer has met with a flower
in which the uppermost sepal was entirely wanting,
while two of the lateral petals were fused together.
Moquin records that in some of the flowers of Cheno-
podiacee, in which the inflorescence is dense, a suppres-
sion of two or three sepals sometimes occurs. The
species mentioned are Ambrina ambrosiodes, Chenopodium
glaucum, 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
Caryophyllacee imperfection as regards the numerical —
symmetry of the flower is not uncommon, as in species
of Cerastiwm, Sagina, Dianthus, &e. In Ranunculacece
the petals are likewise not unfrequently partially or
wholly suppressed. A familiar illustration of this is
afforded by Ranunculus auricomus, in which it is the
exception to find the corolla perfect." Some varieties
of Corchorus acutangulus 1 west tropical Africa are
likewise subject to the same peculiarity. Amongst
Papilionacee absence of the carima or of the alee is not
uncommon, as in Tvifoliwm repens, Faba vulgaris, &e.

Moquin relates a case of the kind in the haricot
bean, in which the cara was entirely absent, and
another in the pea, where both carina and al 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 De Rochebrune, ‘ Bull. Soc. Bot. Fr.,’ ix, p. 281. The author points
out seven grades between complete absence of petals and their presence
in the normal number in this plant. See also Gaudin, in ‘Koch. FI.
Helv. ;? Koch. ‘Synops. Fl. Germ. ;? Cramer, ‘ Bildungsabweich,’ p. 85.

398 MEIOPHYLLY OF THE

In Orchidacee 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 O. arani-
fera; also in Dendrobium mobile, Airides odoratum,
Cypripedium villoswm, Listera ovata, &c. Morren’
mentions analogous deficiencies in Zygopetalum maail-
lave, Oalanthe 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-yeAapov. Mr. Mogeridge
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 andrecium—Suppression of one or
more stamens, independently of lke defects in other
whorls, is not uncommon, even as a normal occurrence,
e.g. m 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 Diplotavis temufolia,? St.
Hilaire in Cardamine hirsuta, others in C. sylvatica.

In Caryophyllacee suppression of one or more sta-
mens has been observed in Mollugo cerviana, Arenaria
tetraquetra, Cerastium, &c.’ Among violets the writer
has observed numerous flowers in which two or three
stamens were suppressed. Chatin* alludes to a similar
reduction in Tropeolum, while in flowers that are
usually didynamous absence of two or more of the
stamens is not unfrequent, ¢. g.in Antirrhinwn, Digitalis,

1 «Bull. Acad. Belg.,’ t. xix, part 1, p. 255.
PEN Ob; 1, p(s tab. 1.0.97.

3 See Gay, ‘Ann. Se. Nat.,’ m1, p. 27.
4 * Ann. Se. Nat.,’ 4 ser., v. p. 305,

.

GYNGCIUM. 399

while in a flower of Cuta/pa a solitary perfect stamen, and
a complete absence of the sterile ones usually present,
have been observed. This might have been anticipated
from the frequent deficiencies in the staminal whorl in
these plants under what are considered to be normal
conditions. Reduction of the staminal whorl is also
not unfrequent in Trifoliwm repens and 1’. hybridwin, and
has been seen in Delphiniwm, &e.'

Meiophylly of the gynecium.—Numerical inequality in
the case of the pistil, as compared with the other
whorls of the flower, is of such common occurrence,
under ordinary circumstances, that in some text-books
it is looked on as the normal condition, and a flower
which is isomerous in the outer whorls is by some
writers not considered numerically irregular if the
number of the carpels does not coincide with that of
the other organs.

But in this place it is only necessary to allude to 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 :—Arenaria 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
im the allied genus Mozinna. Reseda luteola occasion-
ally occurs with two carpels only, while Aconites, Del-
phiniums, Nigellas, and Peeonies frequently experience
a like diminution in their pistil.

In a flower of Papaver Rhwas the writer has recently
met with an ovary with four stigmas and four parietal
placentee only, and to Mr. Worthington Smith he is
indebted for sketches of crocus blooms with two, and
in one instance only a solitary carpel.

Moquin cites the fruit of a wild bramble (/twhus)

' Cramer, ‘ Bildungsabweich,’ p. 90.

4.00 MBEIOPHYLLY.

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 Cvatwgus the pistil 1s
similarly reduced to a single carpel, as in C. monogynd.

The writer has on more than one occasion met with
walnuts (Juglans) with a single valve and a single
suture. If the ovary of Juglans normally consisted
of two valvate carpels, the instances just alluded to
might possibly be explained by the suppression of one
carpel, but the ovary in Juglans is at first one-celled
according to M. Casimir de Candolle.

Among monocotyledons Convallaria majalis may be
mentioned as very liable to suffer diminution in the
number of its carpels, either separately or in association
with other changes.”

Meiophylly of the flower as a whole—In the preceding
sections a reduction in the parts of each imdividual
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 Cobea 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
Talium lancifolium and L. auratwm the writer has fre-
quently met with pentamerous flowers. In Convallaria
maialis a like deviation not unfrequently occurs.’ M.
Delavaud has recorded a similar occurrence in a tulip."

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.,’ xiti, 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 «Bull. Soc. Bot. Fr.,’ viii, p. 287.

MEIOPHYLLY. 40]

four stamens were united by their filaments so as to
form two pairs.

M. Fournier mentions something of the same kind
in the flower of an J7is.’

Orchids seem peculiarly lable to the decrease in the
number of their floral organs. Prillieux? 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 Mpidendrum Stamfordianum. In one
flower of the last-named species the perianth consisted
of one sepal only, and one lip-like petal placed opposite
to it... Morren* describes a flower of Cypripediwmn 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” (Cypripediwn
candidum) “bears two flowers: the axillary one is
normal; the terminal one exhibits the following pecu-
harities. 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, 7.e. on the side of the
style where the labellum belongs. Accordingly the
first impression would be that the labellum is here
transformed into a sterile stamen. The latter, however,
agrees with the normal sterile stamen in its insertion
as well as in shape, being equally adnate to the base of
the style. Moreover, the anteposed sepal is exactly
like the other, has a good midrib and an entire point.
As the two sterile stamens are anteposed to the two
sepals, so are the two fertile stamens to the two petals,
and the latter are adnate to the style a little higher than
the former. The style is longer than usual, is straight
and erect; the broad, disciform stigma therefore

' «Bull. Soe. Bot. Fr.,’ vol. viii, 1861, p. 152. 2 Thid.; ix, p. 275.
* Thid,, 1861, vol. viii, p. 149. * * Lobelia,’ p. 58
26

AO2 MEIOTAXY IN

faces upwards; it is oval and symmetrical, and a light
eroove across its middle shows it to be dimerous. The
placenta, accordingly, are only two. The groove on
the stigma and the placentze are in line with the fertile
stamens.

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

Fic. 198.—Regular dimerous Fie. 199.—Regular dimerous
flower of Calanthe vestita. flower of Odontoglossum Alewandre.

two petals at right angles to the two sepals. The hp
was entirely wanting, but the column and ovary were

THE COROLLA. 4.03

in their usual condition. In Odontoglosswm Alerandre.
a similar reduction of parts has 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 labellum,
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 Melenia was established on a malformed
flower of Orchis of similar character to those above
mentioned.

Meiotaxy of the calyx.—As already mentioned, this
term is here employed to denote those illustrations in
which entire whorls are suppressed. Complete 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 Salicinee 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 per-
foliata and Ruellia clandestina by Linné, who calls such
blooms flores mutilati.' 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 Malpighiacee,
Violacee, Ovalidacee, in some of the flowers of which
the petals are altogether wanting, while in others the

* «Phil. Bot.,”p. 119.

AOA, MBIOTAXY IN

corolla is developed as usual. his 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 Ozalis Acetosella
there are two forms of flower, the one with, the other
without, petals, but both seem equally fertile. Linné
remarks that many plants which, in warm latitudes,
produce a corolla, do not do so when grown in colder
chmates. Thus, certain species of Helianthemwm are
apetalous in Lapland. In the Pyrenees, according to
Bentham, the flowers of Ajuga wa are constantly de-
prived of their corolla.'

Apetalous flowers have been noted most frequently
in the following plants :

Aconitum, sp. pl. ! Crategus !
Cardamine impatiens. Medicago lupulina.
Cheiranthus Cheiri! Melilotus officinalis.
Viola odorata! Ononis minutissima.
Cerastium vulgatum ! Saxifraga longifolia.
Alsine media. Verbascum Thapsus.
Stellaria. Ajuga iva.
Lychnis dioica ! Teucrium Botrys.
Dianthus barbatus, and other Lamium purpureum !
Caryophyllez. amplexicaule.
Helianthemum, sp. ! Polemonium cxruleum,
Oxalis Acetosella. Campanula, sp. pl.!
Balsaminez. Ruellia clandestina.
Malpighiacez. 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, 354. Torrey, ‘Fl. New
York,’ i, p. 428. Hooker and Thomson, ‘Journ. Linn. Soe.,’ ui, p. 7,
Guillemin, ‘Archiv de Botan.,’ i, p. 412. Michalet, ‘Bull. Soc. Bot.
Fr.,’ vii, p. 465. Miller, ‘ Bot. Zeit.,’ 1857, p. 729. ‘ Natural History
Review,’ July, 1862, p. 235.

i

1 *Cat. Plant. Pyr,’ p. 58.

THE ANDRUCIUM. AOD

Meiotaxy of the andrecium—Complete suppression of
the stamens occurs normally in the female flowers of
unisexual plants, and, as an accidental occurrence, is
not very uncommon. Iiricw Tetraliw is one of the
plants in which this is said to happen. ‘The variety
anandra is said to have been known in France since
1635. Cornuti speaks of it in his ‘ Enchiridion.’ In
1860 M. du Parquet discovered it in peaty woods near
Nangis (Seine et Marne).

Many Umbelliferce, such as Trinia vulgaris, present a
like deficiency, while it is of common occurrence
among ftosacew and Pomacee. 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 apply pollen from another flower, a proceeding
made the occasion of festivity and rejoicing by the
villagers in some parts of France. In some of the
Artemisias, especially in Artemisia Towrnefortiana, 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.’
Mr. Moggridge has communicated to the author flowers
of Thymus Serpyllum from a plant in which all the
stamens were deficient, the flower being otherwise
normal,

M. Dupont has given a list of nineteen species of
Chenopodiacece in which female flowers are occasionally
produced, owing to the entire suppression of the
staminal whorl.

Flowers the subjects either of regular or irregular
peloria, g. v., are often destitute of some or all their
stamens, ¢. 7. Calceolaria, Linaria, Viola, &e., 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 andrceecium as a teratological
occurrence has been most frequently noticed in the

' Moquin-Tandon, loc. cit., p. 328.
? For other instances see Chatin in ‘ Ann, Se. Nat.,’ 4 ser., vol. v, p. 305

406 MEIOTAXY OF THE

following plants, omittmg 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! ~ Trifolium hybridum.
auricomus ! repens.
bulbosus ! Umbellifere, sp. pl.

Crucifere, sp. pl. Onagracee, sp. pl.

Violacez, sp. pl. Hippuris vulgaris.

Honckenya peploides. Callitriche vernalis.

Stellaria. autumnalis.

Caryophyllacez, sp. pl. Lonicera Periclymenum.

Malpighiacee, sp. pl. Erica Tetralix.

Tropxolum majus ! Thymus Serpyllum.

Fragaria vesca ! Calceolaria.

Rubus, sp. Composite, sp. pl.

Pyrus Malus. Chenopodiacez, sp. pl.

Agrimonia vulgaris. Stratiotes aloides.

Rosacew, sp. pl.

Meiotaxy of the gynecium—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 Umbellifere, 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 Torenia scabra, accom-
panied by resorption or disappearance of some parts

‘YNG@ICIUM. 4.07

and spiral torsion of others. The pistil was entirely
absent in this instance.’

_M. Gaetano Licapoli places on record an instance
where the petals and carpels of Melianthus major were
suppressed.”

On the whole, the pistil seems less subject to changes
of this character than the andrcecium.

Suppression of the pistil has been most frequently
recorded in flowers (normally bisexual) of—

Ranunculus ! Trifolium repens.
Aconitum ! hybridum.
Delphinium ! Composite, sp. pl.
Peonia. Datura.

Caryophyllee ! Torenia asiatica.
Umbellifere. Colchicum autumuale.

Suppression of ovules,—abortion of seeds.—l'he two cases
are taken together, as the effects are similar, though
it must be remembered that in the one case the ovules
at any rate have been formed, but their development
has been arrested, while mm 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 im other than their native climates
either produce no fruit at all, or the fruits that are
produced are destitute of seed, e. g. Musa, Artocarpus,
&c. Some of the cultivated varieties of the grape
and of the berberry produce no seeds.

Suppression or abortion of the seed is frequently
associated with the excessive development either in
size or number of other portions of the plant, or with
an altered condition, as when carpels become foliaceous
and their margins detached. Hybridisation and cross
fertilisation are also well-known agents in diminishing
the number and size of seeds.

? See also Morren, ‘ Bull. Acad. Bele. xv, Fuchsia, p. 67.
* Cited in ‘ Bull. Soc. Bot., France,’ t. xiv (** Rev. Bibl.”), p. 253.

408 MEIOTAXY.

Meiotaxy of the parts of the flower in general—In the pre-
ceding sections suppression has been considered as it
affected individual members of a whorl or separate
whorls. It rarely happens, however, that the suppres-
sion 18 limited in this way. More generally several of
the parts of the flower are simultaneously affected in
the same manner.

A few illustrations are all that is necessary to give
as to this point.

One of the most familiar instances is that of the
cauliflower or broccoli, where the common flower-stalk
is inordinately thickened and fleshy, while the corolla
and inner parts of the flower are usually entirely
suppressed ; the four sepals can, however, generally be
detected.

Maximowiez describes a Stellaria (Kraschenikovia)
in which the upper flowers are male only, while the
lower ones, which ultimately become buried in the
soil, have neither petals, stamens, nor styles, but the
walls of the capsule are fleshy, and enclose numerous
seeds.'

Karschleger’ mentions a variety of Lonicera Capri-
jolium, 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, xevoc, 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. Flor. Amurens,’ p. 57,
* Flora,’ 1848, p. 484.

SUPPRESSION. £09

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-hy acinth, Hyacinthus comosus,
in which the flowers are almost entirely suppressed,

while the pedicels are inordinately increased in number,
and their colour heightened. Something similar occurs
in several allied species, and in Bowiea volubilis. The
wig plant (hus Cotinus) 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
usual,

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

410 SUPPRESSION.

following :—Ranunculacee, Cruciferae, Caryophyllacee,
Violacee, Leguminose, Onagracee, Jasminacee, Orchi-
dacee. It will be observed that these are all orders
wherein suppression of the whole or part of the outer
floral whorls takes place in certain genera as a constant
occurrence.

Again, it may be remarked that many of these orders
show a tendency towards a regular diminution of the
assumed normal number of their parts; thus, among
Onagracee, Circeia and Lopezia may be referred to,
the former normally dimerous, the latter havmg 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 1s
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 Violacew, Lequminose, and Orchidacee.
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 im others.

As to the suppressions that occur in the case of
the sexual organs, and the relations they bear to di-
morphism, diclinism, &e., 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

SUPPRESSION, 41]

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, &e.

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DEVIATIONS FROM THE ORDINARY SIZE AND
CONSISTENCE OF ORGANS.

Ty 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
detected.

When there is a general reduction in size throughout
all the organs of a plant, or throughout all the nutritive
organs, stem, leaves, &c., and the several portions 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

414, DEVIATIONS FROM

whole of one set of organs be increased in size beyond
the recognised average, we have large varieties, often
qualified by such terms as macrophylla, longifolia,
macrantha, &c. &e. In all these cases either the entire
plant or whole series of organs are alike increased or
diminished beyond ayerage 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 from the
agency of certain conditions promoting or checking
growth, as the case may be; but whether or no, it is
certain that these variations often persist under different
conditions, and that they often retain their distinctive
characters side by side with plants presenting the
normal average dimensions. In other cases the 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 result of

NATURAL SIZE. 415

increased growth and those which arise from diminished
action. It will be 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 are here taken
together. It will be borne in mind that the changes
of consistence from membranous to succulent or woody
are very frequent in the ordinary course of 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.

PAA,

HY PERRO) PAE ay.

Tr 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 fohar 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: i in its usual, healthy, well-formed
state.

It will be seen that under this general head of
hypertrophy, 1 increase of size, homers brought about,
is included; thus, not only increase in leneth, but also
im thickness ; alterations of substance or “consistence,
no less than of dimensions, are here grouped together.

ILYPERTROPHY. 417

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 shght 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 imordinate growth of
some parts is most generally attended by deficiency in
the size and number of others, as in the peripheral
florets of Viburnum or Hydrangea, where the corollas
are relatively very large, and the stamens and pistils
abortive.

CHAP TH Re F

ENLARGEMENT.

A sWOLLEN or thickened condition (renjlement) 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 Hchinocactus, Adeniwm obesum,
some species of Vitis, &e. So, too, the upper portion
of the flower-stalk occasionally becomes much dilated,

27

418 HYPERTROPHY.

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 les beyond the scope
of the present work.

Enlargement of axile organs—All the species of Pe-
largoniwm, Geranium, Mirabilis, as well as those of

Fic. 200.— Pelargonium, one branch of which was hypertrophied.

Caryophyllee and other orders, have tumid nodes as
a normal occurrence. In the genus Pelargonium this
swelling is sometimes not confined to,the nodes, but

KNAURS. 4.19

extends to the interspaces between them, e.g. P.
spinosum. This condition, which happens as a natural
feature im 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 Pelargoniuin
which was thus thickened, the remaining branches not
being in any way affected. The leaves on the swollen
branch were smaller than the others, and their stalks
more flattened. There was, in this instance, no trace
of fungus or insect to account for the swelling of a
single branch, which might, therefore, be due to bud-
variation, perhaps to reversion to some ancestral form.
The repeated cross fertilisations to which Pelargoniums
have been subjected render this hypothesis not an
improbable one.

As an accompaniment to a spiral torsion of the
woody fibres, this distension of the stem is frequently
met with, as in Valeriana, Dipsacus, &c. (See Spiral
Torsion.)

Knaurs.—On certain trees, such as the oak, the horn-
beam, some species of Crategus, &c., hard woody
lumps may occasionally be seen projecting, varying
oreatly 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. rom 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.’ 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

' On the subject of knaurs, the reader is referred to Trécul, ‘Ann.
Se. Nat., 3 ser., vol. xx, p. 65; Lindley, ‘Theory of Horticulture;’ Rey.
M. J. Berkeley, ‘ Gardeners’ Chronicle,’ 1855, p. 756.

4.20 HYPERTROPHY.

the ‘‘uovoli of the olive” and the ‘‘ burrs” that are
formed on some varieties of apple, from which both
roots and leaf-shoots are produced in abundance.

A distinction must. be drawn between those instances
in which the swelling is solid throughout from the
excessive formation of cellular tissue, and those wherein
it is hollow from the more rapid growth of the outer
as contrasted with the mner 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

Fie. 201.—Formation of tubers or hypertrophied buds in the anil of
leaves in the potato.

HYPERTROPHY. 4.21

becomes apparent. A swollen bud or bulbil in this
situation is not uncommon in some cultivated tulips
and lilies. ‘The presence of small tubers in the axils
of the leaves in the potato, as shown in fig. 201, is also
not unfrequent.

Enlargement of the flower-stalk—The cauliflower and
broccoli afford familiar illustrations of hypertrophy of
the flower-stalk, accompanied by a_ corresponding

Fia. 202.—Inflorescence of ash (Firaxinus), with hypertrophied
pedicels, flowers absent.

422, ILYPERTROPHY.

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.’

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.

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Fic. 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. la i,

PEDUNCLE. 423

(See fig. 176, p. 327.) The lower swelling is entirely
axial in these cases, as no trace of carpels is to be seen.
M. Carriere’ mentions an instance wherein from the
base of one apple projected a second smaller one, desti-

Fia. 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 ight, for the reasons already given in the chapter
relating to that subject. (See p. 135.)’

1 «Revue Horticole,’ 1868, p. 110, figs. 12, 13.
* The reader may also refer for further information on the subject of
malformed pears to Irmisch, ‘ Flora,’ 1858, p. 38, tab,i; Lindley, ‘Theory

ADA HYPERTROPHY OF THE

A very curious illustration of hypertrophy of the
flower-stalk is recorded and figured by M. Carriere’ 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.”

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.’

In this place may also be mentioned the hypertro-
phied condition of the placenta observed by Alphonse
de Candolle in a species of Solanwm, 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.* A similar state of things m Lobelia
and Ouphea has already been alluded to under the head
of Alterations of Direction (p. 210).

The following singular growth in a tomato is de-

of Horticulture’ ; Caspary, ‘Bull. Soc. Bot. France,’ vol. vi, 1859 (Rev.
Bibl.), p. 235; Duhamel, ‘Phys. Arbr.,’ liv. i, cap. 3, p. 393, fig. 308;
Bonnet, ‘Recherch. Us. feuilles,’ tab. XXvl, ‘fig. 2; Moquin-Tandon,
‘El. Ter. Veg.,’ p. 384, Xe. 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 Rosacee. The case would fitly be included under altera-
tions of position, but the sheets relating to that subject were printed
off before the pupae of M. Carriére’s notice.

% * Bull. Soc. Bot. France,’ 1860, vol. vii, p. 881.

4 “ Monstr. Veget.,” in ‘ Neue Denkschrift.’

PLACENTA. 4.95

scribed by the Rev. M. J. Berkeley in the ‘ Gardeners’
Chronicle’ for 1866, p. 1217, and appears to have been
an extension of the placenta :—‘ On the first glance it
seemed as if an 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 ines long, and much narrowed below,
besides which, though it was pale green above, the
base was coral-red, hke 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
lacunee, 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 shghtest appearance of placentz 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

426 HYPERTROPHY OF THE

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 oveater 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
merease 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 dise of the leaf, rather than
the margin, 1s increased and its surface 1s 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.

LEAVES, ETC. 427

Such cases as this hardly come under the head of mal-
formations. But where one part only of the leaf
is excessively developed, the other portion remaining
in its ordinary condition, there can be no hesitation in
ranking the phenomenon as teratological.

Thus, Moquin says that the median nerve may be
prolonged beyond the blade of the leaf in the form of
a short strap or ribbon-like excrescence, while, at other
times, the lateral parts of the leaf are subjected to
undue development. He refers to a case cited by
Schlotterbec’ in which each side of the leaves of a
yellow ‘‘ violier”’ (wallflower) was dilated into a kind of
projecting lobe on either side of the true apex of the
leaf, thus rendering it in appearance three-lobed. M.
Delavaud’ 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
jfuscata, which may sometimes be found with small but
perfect leaves projecting from them, the leaf in this
case being the lamina which is ordinarily abortive,
while the scales are the representatives of the stipules.
This condition is said by Hooker and Thomson (‘ Flora
Indica,’ p. 73) to be constant in Magnolia Campbelli.

1 ©“ Sched. de Monst. Plant,” in ‘ Act. Helvet.,’ t. 11, pl. u, f. 14.
2 * Bull. Soc. Bot. France,’ vol. viii, 1861, p. 144,

428 HYPERTROPHY OF THE

Enlargement of the perianth, &e—One or all the segments
of the perianth may be subjected to hypertrophy ; thus,
the utricle of Carex vulpina may frequently be observed
to attain four or five times its usual size, the contaimed
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 Crucifere. The perianth of Iwmex 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 haying found flowers of Salsola Kali
and of Chenopodium murale im 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

Fic. 205.—Hypertrophy of the perianth in Cocos nucifera.

FLORAL ORGANS. 429

appearance of which is due apparently to an hypertro-
phied condition of the segments of the perianth, which
have not only increased in length as the central nut
has ripened, but have developed in their tissues that
fibrous tissue which ordinarily is found in the pericarp
only. This view of the structure of these nuts is
borne out by the fact that, under normal circumstances,
the base of the perianth contains a considerable amount
of fibrous material. In the present case this has 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 Cin-
chonacee, e.g. Mussenda, Pinckneya, Calycophyllum,
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—Polygalacee, Lequminose, Labiate, &e. The
flowers of a rose are mentioned by Moquin as haying
presented an enlargement of the calyx without any
other alterations in form. Schlechtendal has noticed
the same thing in Papaver Rhaas, Reichenbach in
Campanula persicifolia, and A. de Candolle in C. Ra-
punculus. M. Brongniart also has recorded’ a remark-
able variety of Primula sinensis cultivated in the Jardin
des Plantes at Paris, wherein the calyx is enormously
developed. MM. Fournier and Bonnet have described
flowers of Iubus with hypertrophied calyx in conjunc-
tion with atrophy and virescence of the petals and
other changes.”

The corolla may be hypertrophied in some cases,
though the change is more rare than in most other
organs. Moquin-Tandon mentions as subject to this
anomaly species of Galeopsis, Prunella, Scabiosa, and
Dipsacus, and also mentions a remarkable variety of
Viola odorata cultivated in the neighbourhood of

1 «Ann. Se. Nat.,’ ser. 2, t. i, p. 308, pl. ixe, fig. 1.
> Bull. Soc. Bot. France,’ 1862, t. ix, p. 37.

A430 HYPERTROPHY OF THE

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 andrecium—Dunal* 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 Solanwm Duleamara in which
all the upper flowers had two or three stamens of larger
dimensions than the others. This happens habitually
in Solanum tridynamum and S. Amazonicum, and to a
less extent in S. vespertilio and S. cornutum; also m
some species of Hyoscyamus. ‘These cases show the
close affinity between the Solanacee and the Scrophu-
lariacee.

Enlargement of the gynecium.—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 Sueda 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 [vis (Goethe).
MM. Seringe and Heyland’ have figured some anomalous
flowers of Diplotawis tenwifolia 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.
Klinsman’ mentions an instance of a similar kind com-

1 « Gonsid. org. Fleur.,’ Montpell., 1829, 25, 26, pl. ii, f. 18 and 19.
2-* Bull. Bot.,’ t.4, p.°7, tab. 1.
3 * Linnea,’ vol. x, p. 604, tab. 5.

FLORAL ORGANS. 431

bined with hypertrophy of the sepals and pistils; in-
deed, the alteration is not uncommon among Crucifers.
Pyrethrum trodorum 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 hypertrophy
affects also the styles of the central florets, and these
also become enlarged to double or treble their usual
dimensions.

Linne has remarked that the ovary of Tragopogon
sometimes assumes very large dimensions, as also does
the pappus. He mentions a double-flowered variety,
the ovaries of which become ten or twelve times larger
than ordinary. M. Clos’ records an instance in Rumex
scutatus wherein the pistil was hypertrophied or club-
shaped, and open at the top, or in other cases funnel-
shaped, three-lobed at the summit, each lobe terminated
by a style. One of the most frequent causes tending.
to the hypertrophy of the pistil 1s attributable to the
puncture of insects; thus, when the ovary of Juncus
articulatus is thus punctured, it acquires a size two or
three times larger than ordinary, becoming at the
same time sterile.”

Occasionally the enlargement may be due to a fusion
or incorporation of other elements; thus, M. Lemaire
describes an instance in which the style of Sinningia
purpurea was much larger than ordinary, tubular,
bearing three small lobes, and altogether bearing much
resemblance to the column or ‘‘gynosteme”’ of Orchids.
This appearance was due to the cohesion and intimate
union of the styles with three abortive stamens.®

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 *Mém. Acad. Se. Toulouse,’ 5 ser., vol. iii.

> *Ré. nosol. Végét.,’ pp. 342.
* *Tllustr. Hortic.,’ 1868, Misc., p. 62.

432 HYPERTROPHY.

of pea with pods double the ordinary size.’ M. Clos”
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 Leguminose, as Medicago lupulina, Meli-
lotus leucantha, the carpels are sometimes hypertrophied
and elongated, so as to resemble a claw or hook.’

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 Crinum, Amaryllis, and Agave.
These are true seeds enormously dilated.* 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
pomt 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
rearing.

Alterations of consistence often accompany changes im
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 *Mém. Acad. Toulouse,’ t. 6, 1862.

3 *T). C. Prod.,’ ii, pp. 172, 187.

* Richard, “ Obs. sur les bulbilles des Crinum ;” ‘Ann. Se. Nat.,’ t. ii,
p02) pl. a) viel; 2c see also A, Braun, “ Mémoire sur les graines
charnues des “Amaryllidées,” &e.; ‘Ann. Sc. Nat.,’ 1860, vol. xiv, p. 1,
tab. 1.

a

ELONGATION, 433

speaking, the appearances presented in such cases are
not sufficiently striking to demand notice other than
as regards their size. “One illustration, however, may
be cited from its smgularity. This was the case of a
dahha, 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 eemiteined.

CLAP Phi ihr

ELONGATION.

THe class of cases coming under this head are
sufficiently indicated by the name. There are many
instances of this phenomenon occurring under different
conditions, which, though unusual, can hardly be called
abnormal, such, for instance, as the great lengthening
of roots in their search for water, the excessive elon-
gation that takes place in plants when grown at a
distance from the hght, 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 in running water their roots, stems, and some-
times their leaves, become excessively elongated, as 1
Ranunculus jluitans, the flower-stalks of Valisneria
spiralis, &e. These are cases of variation rather than
of malformation, but are none the less curious, or
sometimes perplexing; thus, Lapeyrouse described, m

ABA ELONGATION.

his ‘Supplement a la flore des Pyrenées,’ p. 27, under
the name Potamogeton bifolium, a plant which Mr.
Bentham subsequently discovered to be nothing but a
flowerless variety of Vicia Faba distorted by its growth
in water.’

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
Umbellifere 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 Leguininose, Trifolium repens,
&e. &e.

Another illustration of the sort is that recorded by
M. Fournier, wherein the usually umbellate inflorescence
of Pelargonium was, through the lengthening of the
main stalk, transformed into a raceme. Among Com-
posites a similar change may sometimes be met with.

'* Cat. Plant.,’ Lang., p. 113,

PLOWER-STALKS. AB5

MM. Clos and De Schoénefeld have recorded the ex-
istence of a variety of the sweet chestnut (Castanea) in
which the female catkins were as long, and bore nearly
as many flowers, as the male spikes. This is stated to
be of constant occurrence in some localities, and to
be accompanied by a diminished size of the fruits. A
similar elongation has been observed in the case of the
walnut, catkins of which have been seen bearing thirty
to thirty -five large nuts.’

In the strobile of the hop, Humulus Lupulus, a like
elongation may sometimes be met with, generally in
association with a more or less leafy condition of some
of the scales.

Of a similar character, but complicated with extru-
sion or eversion of an ordinarily concave axis, is the
fig described by Zuccarini,’ and from the appearances
presented by which that author draws the inference
that the peculiar appearance of the fig is due to the
formation of a large number of small bracts blended
together for the greater part of their length, and 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 Amentacee. (See p. 204, figs. 105, 106. )

A lengthening of the axis of the female strobiles of
Conifere is not of infrequent occurrence in Cryptomei “Ld
japonica, Larix europea, &e., and this is usually asso-
ciated, as has been before stated, with a leaf-hke 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 elongation of the main rachis has
been considered. A corresponding deviation occurs in

* Bull. Soc. Bot. France,’ t. i, 1854, p. 175, and t. xi. p. 96.
‘ Abhandl. Math. Phys. Class..’ Band. iv, Abhandl. i, fal i.

436 ELONGATION OF THE

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-

Fic. 206.—Inflorescence of Ranunculus acris, with secondary peduncles
lengthened.

presents a specimen of Ranunculus acris, 1 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 repens, Lotus cori-
culatus, &c., what is usually a compact spike, or head

LEAVES. AS7

of flowers, becomes a raceme from the elongation of
the pediceis. In Umbellifere 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 Razoulzi, Carwm Carwi,
Thysselinum palustre." In Composite, when affected
by an analogous change, the capitulum assumes the
appearance of a simple umbel, as in Hypocheris radicata,
Senecio vulgaris, 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 imstances 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, e.g. Ranunculus fluitans, and in terrestrial
plants there are varieties termed longifohar, from the
unusual leneth of the leaves. <A similar lengthening
occurs in the involucral leaves of Umbellifere and Com-
posite, changing very materially the general aspect of
the inflorescence. Occasionally, also, the leaf-lobes of
parsley (Apium Petroselinum) and other crested-leaved
plants may be observed to lose their ordinary wavy
form, and to be lengthened into flat riband-lhke 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 Culturpflanzen.’

438 ELONGATION OF

the lobes or leaflets become separated one from another
bya kind of apostasis. This change may be frequently
seen in the horse-chestnut, particularly in the young

Fra. 207.—Portion of leaf of parsley, showing the change from short
wavy to long flat leaf-lobes.

shoots formed after the trees have been pruned or
pollarded. In the adjoining cut the intermediate stages
between a palmate or digitate leaf to a pinnate one
may be seen. The specimens from which the drawing
was made were taken from the same tree at the same
time.
In the white clover, Trifolium vepens, a similar tran-
sition may often be observed, as also in some species
of Potentilla.'

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; ‘Linnea,’ xi. p. 301, xiv,
p. 363 ; ‘Bot. Zeit.,’ 1856, p. 72 ; Masters, ‘ Rep. Brit. Assoc.,’ Manchester,
1861; Coultas, ‘What may be learnt from a tree,’ p. 11S.

FLORAL ORGANS. 439

the carpels, sometimes as a result of injury from in-
sects or fungus, at other times without assignable
cause.

Fie. 208.—lLeaves of horse-chestnut, Hseulus, showing passage from
“digitate to pinnate leaves.

In the case of inferior ovaries this lengthening 1s,
perhaps, even more common, as in Umbellifere, Com-
posite, &e. The common groundsel (Senecio vulgaris)
is especially liable to this form of enlargement of the

440 APOSTASIS.

pistil, either im association with a leafy condition of
the pappus or without any such change.

Elongation of the thalamus, placenta, &e—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-bearmeg shoot.’

Apostasis—Hngelmann made use of this term to ex-
press the separ ation of parts one from another. by the
unusual elongation of the internodes.” 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, ‘&e. ., of Passi-
floree, Caryophyllee, Capparidee, &c. &e.

A similar elongation of the thalamus, bringing about
the separation of the floral whorls, or of their con-
stituent parts, 1s 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 «De Antholysi,’ p. 42, § 49.

APOSTASIS. 441

and are, in some cases, united by their margins, so that
a spiral sheet or tube is formed, surrounding the axis.
This frequently occurs in double flowers of the Chinese
primrose, Primula sinensis.

Engelmann’ figures a case wherein the calyx of
Anagallis phenicea was separated by a rather long in-
ternode from the corolla, and a like illustration in
Torilis Anthriscus.

Fic. 209.—Flower of Delphinium, showing apostasis of carpels, from
lengthening of the thalamus, Xc. (Cramer.)

1 Loe. cit., tab. 2, f. 6.

4,49, APOSTASIS.

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 1m
Capparids.

In Tropeolum imajus 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 Hpilobium hir-
sutwm, specimens of which were originally obtained
from the late Professor Henslow. The several floral
parts are here, some virescent, others truly foliaceous,
and each whorl is separated from its neighbour by a
rather long internode. In Fuchsia and Campanula a
hike 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
Chamedrys, Orobanche gracilis, Solanum Lycopersicum,
Gentiana campestris, Hypericum, Helleborus fetidus,
Caltha palustris, Brassica oleracea! and many Rosacee,
Caryophyllee, Crucifere, and Ranunculacee. (See
Dialysis, Median Prolification, &c.)

Apostasis of the subfloral or involucral leaves is not
of infrequent occurrence in malformations affecting
Composite and Umbellifere. In the following genera
it has been observed with especial frequency :—Torilis
Anthriscus, Eryngium, Athamanta Cervaria, Leontodon,
Tragopogon pratense !, Wedelia perfoliata! In garden
anemones, also, it is a common deyiation.- ©

OHA Pare ne PP.
ENATION.

Unper 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-
erowths 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 Caryophyllew, Sapindacee, &e.,
the coronal filaments of passion-flowers, the cup of
Narcissus, the appendages that beset the segments of
the perianth in Laliwm lancifoliwm, and other similar
erowths, may be referred to a lke process. In many
cases this has been proved by a study of the develop-
ment of the flower, from which it appears that the
erowths 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

AAA, ENATION.

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
erowth 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 enaurs 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-
minose, nor of the peculiar excrescences on the roots
of Taxodiuwm distichum, as these appear to be normal
formations. But it may be well to mention im this

ENATION. 4AD

place an anomalous development which occurs occasion-
ally in Ruscus aculeatus, and in which, from the upper
surface of the ordinary flattened leaf-hke branch, pro-
jects at right angles a second similar branch, so that
in section the appearance would be like that of the in-
verted letter t; thus, J.

Enation from foliar organs—Leaves—'l'he 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 lamine projecting at right angles from
the primary one may frequently be observed. In the
instance figured (fig. 210) the new growths proceeded

Fic. 210..—Section through base of midrib of cabbage leaf, showing .
supplementary lamine, Xe.

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,

44.6 ENATION.

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 © . 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.' (See p. 33.)

' Tt is desirable in this place to allude to a singular case of fissiparous
division of a leaf of Prunus Laurocerasus described by Prof. Alexander
Dickson (‘Seemann’s Journ. Botany,’ vol. v, 1867, p. 323), and which
did not come under the writer’s notice till after the sheet relating to
fission, p. 61, had been sent to press. Dr. Dickson thus speaks of this
abnormal leaf :—‘* The petiole (unchanged) supported two laminz, placed
back to back, and united by their midribs (7. e. not separated) to within
about an inch from their extremities, which were perfectly free from
each other. These lamine 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

x
represented : \O O/. The nature of the case may be even better seen

5 C ‘ x

by comparison with the normal arrangement, which would be o .

——"
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:

x

Say , the x in all cases representing the position of the axis, the

O that of the axillary bud, and the —— that of the lamina.

ENATION. 4,4,7

Such leaves as those of the hedgehog holly, [ex
Aquifolium, var. feror, and, to a less extent, bullate
leaves, may also be mentioned here as illustrations of
hypertrophy or enation.

Fie. 211.—Nepkrodiwm 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.”* These

1 Linn., ‘Phil. Bot.,’ § 274. The term “crispa” is surely preferable
to that of Ré, “ phyllorhyseme.”’ a€

448 ENATION,

conditions occur normally in such leaves as those of
Rumez crispus, Malva crispa, &c.,and are developed to an
extreme degree in garden varieties of parsley, some kails,
&e., as well as in many ferns, but these are probably
cases rather of fission than enation as here understood.'

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 mner surface
of the petals, &c. Under natural circumstances the
formation of scales, lobes, &c., from the petals, as in
some Caryophyllee, Sapindacew, &c. &e., may be ex-
plained, as already remarked, by this process, rather
than by fission, chorisis, or by substitution of petals
for stamens, &e. Hach 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 followmg
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 déformations,” &c., in ‘ Bull. Acad.
Belg.,’ 1852, tom. xix, part 3, p. 444; and as to ferns, see Moore,
‘ Nature- Printed British Ferns,’ 8yo ed., where numerous illustrations
are given,

FROM THE COROLLA. 44.9

of alternation) that 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
simensis, 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 —~; while, in other
instances, the front of the adventitious lobe is directed
towards the corresponding surface of the original petal,
thus <>. 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

29

A5O ENATION

already referred to at p.446. In some flowers of Datura
jfastuosa 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

Fie. 212.—Datura fastuosa. True corolla turned back to show the
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
mner corolla were opposite to those of the outer one,
the intermediate petalodes were opposite to both; thus:

S 8 S S S
P P P P P
x x x x xX |
cr au Site |
B P iP 12 Pj

st st st Sb o®- ING

FROM THE COROLLA. 45]

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,! but which is now becoming common in
English gardens. When first observed these flowers

Fic. 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. Being encouraged and tended
by gardeners, in course of time, instead of a series of
petalodes, more or less distinct from one another, a
second corolla or “catacorolla’’ was formed outside
the primary one, so that a hose in hose flower was
produced, but, in this case, the supplementary flower
was formed on the outside and not within the ordinary
corolla. Moreover, the disposition of the colour was
reversed, for in the outermost corolla the richest hues
were on the outer surface, while in the inner or true
corolla they were on the inside.

' «Bull. Acad. Belg.,’ t. xix, p. 224, tab.i; and ‘ Gardeners’ Chronicle,’
1865, p. S65,

452 ENATION

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
question.

—

Fig. 214.—“ Catacorolla” of Glowinia, 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-

FROM THE CARPELS. 453

titious lobes from the outside of the minute partially
developed petals could be distinctly seen.

Enation from the stamens——An illustration of this process
occurred in some double-flowered rhododendrons, which
presented the following arrangement of parts :—calyx
and corolla normal; within the latter eight petal-lke
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
EN

—-~. This disposition was well seen in the anther,

OO

half of which was, in some cases, petaloid like the
filament; in fact, the mner wing of the latter was
directly continuous with the petal-hke expansion from
the anther. A section through the latter showed,
going from within outwards, the cut edges of two
perfect polliniferous lobes in the centre; and on either
side the petaloid wing representing the remaining
anther-lobe ; outside these were the edges of the re-
maining wings, one on each side. (See p. 290, fig. 155.)

Enation from the carpels—-The only imstances 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 writers on the genus Citrus,
may be mentioned under this head. <A similar forma-

tion occurs in the fruit of some species of Solanum.
(See p. 316.)

far iNe tt Le ie
ATROPHY.

Unver 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 shght 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. nane), or those in which the leaves or flowers are
smaller than usual (var. parvifolie, v. parviflore), are
truly regarded as variations, and not as malformations
properly so called.

ABORTION, 459

Atrophy is partial and special in its operation, nanism
is general.

Under ordinary circumstances atrophy is exemplified
by the presence of rudimentary or imperfect organs, as,
for mstance, in Pentstemon, Scrophularia, &c., where
one stamen is atrophied.

For convenience sake atrophy may be divided into
abortion and degeneration, the first including cases
where, from ar rest. of dey elopment. occurring at an
early stage, organs are present; but im 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 re ather perverted, so that not only
the dimensions are lessened, but the form is altered.

CHAPTER I.

ABORTION.

Tux 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 1s
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, plante acaules. In these the inter-

4.56 ABORTION OF THE

nodes are so shehtly developed that the leaves are
closely crowded in tufts or rosettes. When this short-
ening of the stem (acaulosia) occurs, without other
considerable change in other organs, the deviation is
classed under the head of variation rather than of
monstrosity; and, indeed, in very many plants, this
arrested growth of the axis is the rule rather than the
exception. When occurring in an abnormal manner,
atrophy of the stem is most frequently attended by
other more or less grave alterations in other structures ;
thus Moquin-Tandon' cites an instance of Campho-
rosma monspeliaca, wherein the stems presented the
form of very short, hard, woody tubercles, thickly
clothed with deformed leaves, and imvested by a vast
number of hairs, longer and more dense than usual.
A similar deformity sometimes occurs in an Indian
species of Artabotrys ; im 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.”

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.°

1 «Hl. Ter. Veg.,’ p. 182.

2 Spinose arbores cultura sepius deponunt spinas in hortis, ‘Linn. Phil.
Bot.,’ § 272.

3 Mr. Selby, in his ‘History of British Forest Trees,’ p. 465, gives

RECEPTACLE. 457

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 receptacla—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 Composite and Umbellifere. In the former
natural order the following plants have been met
with in this condition: — *Tragopogon pratense !,
*Cirsium arvense, Hypocheris radicata, Senecio vulgaris!,
Coreopsis Drummondi. In the latter order, Daucus
Carota! Cinanthe crocata! and Thysselinwm palustre,
seem most frequently to have been observed in this
state.. 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
lace.”
: 1 See Cramer, ‘ Bildungsabweich.,’ pp. 53, 64, for further references,

458 ABORTION

partially developed only, and forming a kind of cup,
from which the true carpels protrude.

Abortion of the leaves—Arrest of growth in the leaves
occurs in different ways; sometimes the whole leaf is
smaller than usual; at other times certain parts only
are reduced in size; while, in a third class of cases,
portions of the leaf are entirely suppressed.

Moquin' mentions haying seen the leaves of Cheno-
podium vulvaria, and of Diplotazi is muralis reduced to
a fourth of hee natural size; and he alludes to other
cases of the same nature, seen by other observers, in
Hypericum perforatum and Blitum polymorphum.

Nicandra physaloides’ 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 Coronopus, var. integrifolia, Papaver
Rheas integrifolia, &c. &e. The same remark may be
made of those specimens in which one part of the leaf
is developed to a less extent than another, as happens
in the submerged leaves of such plants as Ranunculus
aquatilis, Cabomba aquatica, the spiney leaves of 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 «Hil. Ter. Veg.,’ p. 124.
2 Schlechtendal, ‘ Bot. Zeit.,’ 1857, vol. xv, p. 67.

OF THE LEAVES. 459

thus altered, but the following species appear to be
particularly subject to this change : Asculus Hippocas-
tanum, Digitalis purpurea, Morus alba, Fagus silvatica
contracta (hort.), Codiwum variegatum var. eroswm
(hort.), Broussonettia papyrifera, Scolopendrium vulgare,
&e.

Frequently this irregular diminution in proportion
is coexistent with an unusual degree of cleavage or
laciniation of the margin, as in Acer platanoides lacinia-
tum, Tilia asplenifolia, Alnus imperialis (hort.), Fagus
silvatica var. (hort.), &e.

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 laminez one with the
other. This has been observed amongst other plants
in Veronica latifolia, Broussonettia papyrifera, Codieum
variegatum var. interruptum (hort.), Scolopendrium vul-
gare, &c.’ (See p. 328.)

In some of the leaves which have been already re-
ferred to in illustration of the mordinate growth of
the cellular portions, the mcreased 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, Coleus, Fagus silvatica crispa, Cytisus,
Laburnum var., and other forms, cultivated in gardens
for their singularity.

Entire absence of the stalk of the leaf occurs 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 Ozalis, Indigofera,
Tebeckia, Ranunculus, Bupleurum, &e.

1 On the subject of this paragraph the reader may consult A. Braun,
“Ueber abnorme Blattbildung,” &c., in ‘ Verhandl..’ d. 35, ‘ Naturfor-
scherversammlung ;’ Jaeger, ‘ Flora,’ 1850, p. 481, tab. 4, Digitalis.

4.60 ABORTION

De Candolle,' from a consideration of Strelitzia juncea,
in which the petiole alone is developed, was led to the
inference that in many monocotyledonous plants the
blade of the leaf was never developed, the portion
present being the sheath or stalk, unprovided with
limb. The correctness of this inference is shown,
amongst other things, by the occasional presence of a
leaf-blade in Strelitzia juncea itself.

Occasionally the laminar portions of the leaf are
completely wanting, leaving only the main ribs, as in
the case of Berberis, while the adjoining figure (fig. 215)
represents an instance of a cabbage wherein the inner-

Fig. 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.—Lllustrations
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.

OF PERIANTH, BTC. 461

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 1s more common in the case of the corolla than in

Fie. 216.—Abortion of four out of five petals, Viola tricolor, side and
front views.
that of the calyx. In Arenaria serpyllifolia the petals,
especially in autumn, are only one fourth the length of
the sepals. Anagallis phenicea, Honckenya peploides,
Arabis alpina, Ranunculus auricomus, Rubus fruticosus,
and Geraniwm 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 Violacee.

The diminished size of the petals sometimes coexists
with an increase in their number, as in a flower of
Streptocarpus Rewti, mentioned by Bureau.*

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. 710.

462 ABORTION OF THE

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’ mentions a flower of [ritillaria vm-
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’ records a similar
occurrence in the same plant.

Morren’ gives details of like appearances in Hyimeno-
callis americana, and Delavaud'* in Tigridia pavonia.

In certain orchids an arrested development of the
perianth is habitual, as im Oncidiwm abortivum (fig.
217), where, on a large branching panicle, numerous

Fic. 217.—Flower of Oncidium abortivum, magnified.

abortive, but few perfect, flowers are produced. Ina
similar way the petals and labellum of Odontoglosswm

‘Linnea,’ 1830, vol. v, p. 492.

‘Mus. Senkenb.,’ ii, p. 45.

‘Bull. Acad. Roy. Belg.,’ 1851, t. xviii, part i, p. 275.
‘ Bull. Soe. Bot. France,’ vol. viii, 1861, p. 147,

mw Ww

STAMENS. 4.63

Uro-Skinneri have been found reduced to filamentous
processes.

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. Scrophularia, Hrodiwn,
many festiacew, &c. &e. As a strictly teratological
condition atrophy of the stamens is more rare than
complete suppression. It has been noticed in Arabis
alpina, Cerastiwum glomeratum, C. tetrandrum, Rhamnus
catharticus, Anemone, Hepatica, &e. 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 stili
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.' 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. Anim. and Plants,’ ii, 165.

A464 ABORTION OF THE

atrophied, and distorted, and contained little or no
pollen; the few grains of the latter being smaller than
usual. (See under Heterogamy, pp. 193—196, and
p- 398.)

Abortion of the pistil, fruit, &e.—Traces of the carpels
occur in many male flowers of unisexual plants, e. g.
Sterculiacee, Huphorbiacee, Restiacew, &c. &c., and in
some natural orders there appears to be a tendency
towards a dioecious condition, e.g. Caryophyllee, as in
Iychnis dioica, Silene otites, Arenaria tetraquetra, Xe.
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.’ The
ovary of aconites, according to Moquin, is very subject
to atrophy.

W A 2 Cie,
Ws SS | A
UN

Fic. 218.—Bladder plum.

During the maturation of the pistil, and its passage
to the fruit, great changes of consistence frequently

1 Gay, ‘Ann. Se. Nat.,’ ser. i, 1824, t. ii, p. 44.

PISTIL. 4.65

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.'

MM. Fournier and Bonnet’ describe a fruit of a
Rubus, with perfectly dry fruits, like those of a Gewm,
and this form was considered by Steudel to form a
distmet species. It is, however, merely a variety in
which the fruits have not become succulent.’

Schlechtendal describes* the ordinarily baccate fruit

' See De Candolle, ‘Mem. Legum.,’ tab. 3, f. 1; Wyville Thomson,
‘Trans. Bot. Soc. Edinb.,’ 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.

2 * 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.

+ *Tinnea,’ vol. v, 1830, p. 493.

30

4.66 ABORTION.

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, leay-
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.’ See also under Heterogamy, Meiophylly,

&e.

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.” 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.

ABORTION. 467

or imperfect development of the indusium, as in what
are termed exindusiate varieties.’

General remarks on abortion, coincident changes, &c—Refer-
ence has already been made, while treating of hyper-
trophy, suppression, &c., to certain other changes
affecting the flower at the same time. Atrophy of one
organ or set of organs, for instance, is frequently
accompanied by a compensating hypertrophy or by an
increased number of other parts. In the feather-hya-
cinth, Muscari comosum, var., monstroswm, the absence
of flowers is compensated for by the inordinate for-
mation of brightly coloured threads which appear to
be modified pedicels (see pp. 347, 348); so also in the
wig plant, Rhus Cotinus. So the atrophy of the 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 Desmodium marylandicum were atrophied,
while the calyx and legume, on the other hand, were
hypertrophied.

Fusion of the members of one whorl with one
another, or with the components of an adjacent series,
often entails atrophy or suppression, either in the
united organs themselves, or in adjacent ones. A
fohaceous 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’®
on the different degrees of atrophy up to complete sup-
pression, observable in the flowers of Bellevalia comosa,
are of importance. According to this observer, the
most highly differentiated parts, such as the stigmas,
the ovules, and the anthers, are the first to disappear,

1 See Moore, ‘ Nature-Printed Ferns,’ 8vo, for numerous illustrations
both of depauperate and exindusiate ferns. Scolopendrium vulgare
seems to be one of the ferns most commonly affected im this way.

Moore, loc. cit., vol. ii, pp. 135, 147, 159, 165, &e.
2 Bull. Acad. Belg.,’ t. xvii, p. 38, t. 1; Lobelia, p. 85.

4.68 ABORTION.

the filaments often bemg 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
pistils.

The impotence of the pollen in certain atrophied
flowers, as noticed by the same observer, is of much
interest, especially in reference to the sexual relationship
between the different forms in polymorphic flowers as
studied by Mr. Darwin.

A change in direction may also be noted as a common
accompaniment of atrophy or suppression; thus, in a
capsule of Veronica Beccabunga, which was one-celled
by the abortion of one carpel, the style was lateral
instead of terminal.

As to the causes of these structural deviations butlittle
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’ 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 ;
A, 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.

ABORTION. 4.69

culiar conditions of cultivation; thus, cuttmmgs and
layers produce sterile and abortive seeds much more
frequently than plants of the same species raised from
seed; 6. abortion of the seed is often combined with
luxuriant development of the walls of the fruit.

Temperature and climatal changes in general seem
not to be without effect, as has been already mentioned
in the case of Arenaria tetraquetra, which 1s polygamous
when growing in mountain districts. Other illustra-
tions of a similar character are mentioned under the
head of Heterogamy (p. 196).

Pressure has been already alluded to as one of the
most obvious of the inducing causes of atrophy and
suppression.

In the case of Ranunculus auricomus before cited, in
which the petals are rarely perfect, M. de Rochebrune
considers that the deficiencies in question depend, in
great measure, on the amount of moisture in the
localities where the plant grows. In most places the
flowers and carpels are apt to become more or less
abortive, while the leaves are luxuriant; while, in dry
places, the foliage is small, but the flowers are more
perfect. This is quite consonant with other facts 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 1s almost cer-
tain that internal causes inherent to the organization
of the plant are more important. Mr. Darwin’ 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 dicecious, as in Caryophyl-
lacee, Orchidacee, Plantaginacee, Primulacee and other
orders.

1 «Origin of Species,’ p. 450.

470 DEGENERATION,

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. Veget.,’ p. 120; C. Morren, “De Vatrophie en général,” in
‘Bull. Acad. Belg.,’ t. xviii, 1851, part i, p. 275.

ChE AP a He a ae

DEGENERATION.

Watts 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, 7. 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 perule of leaf-buds, or the pales on the
common receptacle of composite flowers. Other illus-
trations of a like character are to be met with in the

DEGENERATION. ATI

membranous scales that represent leaves in Ruscus,
Asparagus, Pinus, &c. Similar productions are met
with within the flower, where they may occur as the
representatives of sepals, petals, stamens, or pistils, or
as mere excrescences. (See Hnation.) Whole families
of plants, e. g. Sapindacee, are characterised by the
presence of these organs, which are often of great
interest to the morphologist as indicating the true
symmetry of the flower, while they have acquired fresh
importance since the publication of Mr. Darwin’s work
on the ‘ Origin of Species,’ wherein we are taught to
regard these rudiments as, in many cases, vestiges of
organs that were more completely developed in the
progenitors of the present race of plants, and the
exercise of whose functions, from some cause or other,
haying been rendered impossible, the structures become,
in process of time, proportionately stunted.

Thus, in dicecious 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, asin 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.

472 DEGENERATION.

Moquin cites in illustration of the first phenomenon
the flower of a Vicia, 1m 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-
elove 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
uncommon.

Formation of hairs, spines, &—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.

DEGENERATION. 473

They occur normally in the sepals of Paronychia
serpyllifolia and other plants.

Formation of glands——Under this name are associated
a number of (generally) rudimentary organs very
different in their morphological nature and sionificance,
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 Opulus, the so-called glands
on the sides of the petiole appear to represent leaflets,
and are not unfrequently developed as such.

M. Dunal observed a flower of Cistus vaginatus in
which some of the stamens were replaced by an hypo-

eynous disc.’ Moquin has seen similar instances in
the flowers of a Rose, Hypericum, and Poppy.

M. Planchon’ 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 Trécul.’

Dr. Hooker shows that the pitcher of Nepenthes is
due to a modification of a gland placed at the extremity
of the midrib.*

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, pl. ii, fig. 23.

2 «Ann. Sc. Nat.,’ 3 ser., Bot. ix, pl. 6, ff. 1, 2.

3 * Ann. Se. Nat.,’ 3_ser., Bot. 1855, pp. 297, et 303.
4 «Trans. Linn. Soc.,’ xxii, p. 415.

ATA, DEGENERATION.

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.*

Kairschleger, however, has recorded the existence of
a cirrhose sepal in Curcurbita Pepo.’

1 See Darwin, “On Climbing Plants,” ‘Journal of Linnean Society,’

vol. ixpals
2 * Flora,’ 1845, p. 615.

GENERAL CONCLUSIONS.

At the end of many of the preceding sections, and
whenever the requirements of the case demanded it, a
brief summary of the main facts and of the inferences
to be derived from them has been given. It may be
useful to give in conclusion a few general remarks on
the whole subject.

It will be seen from the numerous facts herein cited,
that the so-called monstrous formations (excluding
morbid growths the result of disease or injury) present
no peculiarities absolutely foreign to the normal
organisation of plants. The difference between the
natural and monstrous development is one of degree
and frequency of occurrence, not of kind.

Deviations from the customary form have been
shown to arise from excessive or diminished growth,
or from arrested or exalted development. Hven in
those imstances 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

A76 GENERAL MORPHOLOGY.

erowth 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

MORPHOLOGY. 477

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 the development and construction of the leaf -
further than they are illustrated by ordinary terato-
logical phenomena.

From this point of view perhaps the most interesting
circumstance is the part that the sheath of the leaf
plays.’ 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
erowing point (saillie) and its corresponding fibro-
vascular bundle.*

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.
> «Théorie de la Feuille,’ p. 26.

478 MORPHOLOGY.

thoroughly recognised that little need be said on that
score, save to repeat that the homology of the floral
organs is usually not so much with the entire leaf as
‘ with its sheath.

The most singular instances of morphological identity
are those relating to the sexual organs. We have seen
the gradual transition of stamens to pistils, and of pistils
to stamens, the development of ovules on the edges of
the anther, the co-existence of pollen with ovules on an
antheroid body, and, stranger still, the actual develop-
ment of pollen within the tissues of the ovule itself!
From such facts, in addition to what we know of
the relative position, internal structure, and mode of
development of the organs, it 1s 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, may be advanced in confirmation of the opinion,
that, morphologically, there is no distinction between
axis and leaf. The leaf, according to this view, is a
specialised portion of the axis set apart to do certain
work, just as the petals, stamens, &c., are leaves told
off for distinct uses. It is unnecessary to refer to the
intermediate productions linking the leaf-form to that
of the axis, all that is requisite here is to pomt out |
the facts that teratology lends in support of these views.
These may be summed up by the statement that almost
all those attributes which morphologists recognise as
peculiar to one or the other organ respectively, may
be and are manifested by both. We have the stem
acquiring the characters of the leaf, and the leaf those of
the stem. Thus we have seen leaves, leaf-buds, branches,
and fiower-buds springing from leaves or leaf-organs ;'
see pp. 174, 177, 445, &. The structure that we

1 An additional illustration of this may be cited, which has been

MORPHOLOGY. 479

are apt to associate exclusively with one is found to
pertain tothe other. The arrangement of the vascular
cords in the leaf-organ finds its counterpart in the axis,
generally, it is true, modified to suit altered circum-
stances or diverse purposes. In some cases the dis-
position is absolutely indistinguishable in the two
organs. It may then be said that the distinctions
usually drawn between axis and leaf are not absolute,
and that, however necessary such a separation may
be for descriptive or physiological purposes, 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 Cryptogams,
but which in the higher groups of plants becomes
marked out into separate portions, each portion haying
its own distinct functions to fulfil for the common
benefit of the whole organisation.’

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 recently, and
in which some of the leaflets of the pinnate leaf of a species of Macro-
lobium were absent, and their place supplied by flowers arranged in

es.

} The presence of a bud at the extremity once considered to be an
absolute distinction between branch and leaf, which latter never forms
a bud exactly at the apex—is invalidated by the case of the Nepaul
barley, p. 174.

A480 CALYX-TUBE.

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 imsepa-
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 1s to be
regarded as composed of the confluent bases of the
sepals.

Mr. Bentham, who has recently reviewed the evidence
as to the nature of the calyx-tube in his paper on
Myrtacee,' 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.”

Organs originally free and distinct become ultimately
combined at the base by the gradual protrusion from

1 « Journ. Linn. Soc.,’ vol. x, p. 103 ef seq.

2 See also the receptacular tube (ovary ?) of Baeckea bearing stamens,
see p. 183. It would be natural to see stamens springing from the recep-
tacle but not from the ovary.

OALYX-TUBE. 481

the receptacle of a ring or tube under them, as in the
stamens of Leguminose ; yet, says Mr. Bentham, no
one would propose to describe the staminal tube of
monadelphous Leguminose 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 O47, 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 Rosacew, Myrtacece, Melasto-
mace, Passiflora,' &e.

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.’ 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-lixe 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 Tropwolum or Nelumbium
become very significant. In both the leaf-stalk is
cylindrical and tr aversed, as in the case of all cylin-

‘ In Passiflora the organogeny of the flower clearly shows the truth

of this assertion, as was indeed shown by Payer and Schleiden.
2 See Payer, ‘Organ. Veget.’
[3 )
ol

482, INFERIOR OVARY.

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).

Andrecium.— 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
andrcecium 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 tobe purely axial. On the contrary, the evidence
derived from this source supports the ordinary opinion

PLACENTATION,. 483

that the carpels are invaginated within the expanded
top of the flower-stalk and more or less adherent to it.
Some of the gourds afford good illustrations of this, the
upper part of the carpels in these fruits projecting
beyond the axial portion. But this matter loses much
of its importance if the morphological identity of axis
and leaf-organ be conceded. The carpels in inferior
ovaries seldom or never correspond to the lamina of the
leaf, and between the vaginal portion of the carpellary
leaf, and the axis who shall draw the distinction ?

Placentation—Some botanists have considered the
placentas to be portions of the carpel, and have com-
pared the production of ovules on them to the forma-
tion of buds on the leaf of Bryophyllum. Others
have been led to see in each placenta, even when
it is, to all outward appearance, a portion of the car-
pellary leaf, a direct prolongation from the axis, ad-
herent to the leaf. Teratology shows that ovules may
be formed indifferently on leaf-organs or on stem-organs.
Sutural, parietal, axile, free-central placentation, and, if
there be more forms, all may be met with even in the
same ovary (see pp. 96,508). Now, if there were such
special tendencies in the axis, as contrasted with the
leaf, to produce ovules, it 1s 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,’ there is surely little ground
for considering the placentas, or ovuliferous portions
of the plant, to be of necessity axial. Here again,
much of the difficulty vanishes if the morphological

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 impari-pinnate leaves
in which the terminal leaflet is jointed to the common rachis, and in
the leaves of some Meliacew which continue to push farth new leaflets
even after the leaf has attained maturity.

A484 MORPHOLOGY.

identity of the leaf-form and of the stem-form be ad-
mitted.

Structure of the ovale—The nature of the ovule and of
its coverings has been a fertile source of controversy.
The teratological data bearmg on this subject have been
given at pp. 262—272. These data strongly support
the notion of the fohar nature of the coatings, and of the
axial nature of the nucleus, taking leaf and axis either
in the ordimary 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 Chevranthus, not uncommonly
met with (p. 182), are instances of ovular transmutation
may be open to doubt.

The axial nature of the nucleus has been inferred from
its position, mode of growth, and from its occasionally
lengthening into a leafy or even a floriferous shoot.
Probably it may occasionally be invested by sheathing
coats, more analogous to tubular processes from the
receptacle, than to folar organs, as is the case im
Welwitschia. The discussion of this matter, however,
pertains rather to normal morphology than to tera-
tology.

Morphology of conifers—'T'he nature of the pseudo-leaves
of Sciadopitys, and probably of other Conifers, is illus-
trated by teratolog ey, 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

MORPHOLOGY. 485

development, the persistent union of parts, usually
separate in the adult state, has been traced to an
arrest of the process of development, by no means
necessarily coexistent with diminished growth. The
diminished or increased number of parts is, in like
manner, attributable to analogous causes, as also are
the variations in arrangement and form, spoken of
under the heads of Displacement, Peloria, 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 brig 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 briefiy alluded
to. The deviations from the customary arrangement
have been very generally attributed to suppression,
or to chorisis. It is unquestionable that either of these
affords an efficient explanation of the arrangement in
question, as also does that modification of chorisis, as
it may be considered, which has been treated of under
the head of Enation. Spiral torsion of the axis would
likewise bring about analogous results. Still, it is quite
conceivable that opposition or superposition of organs
may occur without the intervention of any such 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-

486 CO-RELATION.

position .of these cords have also been considered to
bring about deviations from the rule of alternation,
but in general the formation of the cords is subsequent
to that of the growing points or mamelons.

Adhesions, accompanied by displacements, 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.’
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 with perfectly green, long, tubular, erect, not
horizontally spreading flowers. ,

CO-RELATION. 487

There are certain co-relations which do not appear
to have hitherto attracted the attention they merit ;
such, for instance, is that which exists between the
particular period at which an organ is developed
and its position and form. In normal morphology
this has, to some extent, been worked out, as in
the case of definite and indefinite, centrifugal and
centripetal inflorescences, and in the definite or 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, triplicati, &c.), and also with
cases of peloria. In instances where the organs are
formed successively in spiral order, we meet with such
changes as median prolification, petalody, and phyllody.
All these are alterations which we might anticipate from
the activity of the growing point being checked at a
certain stage in the one case, while it is continuous in
the other. This relationship between the definite and
indefinite modes of growth and the form of the several
organs of the flower, is more constant in reality than
it may appear to be from a perusal of the lists of
genera in the foregoing pages, in which it was not
possible to show sufficiently well the comparative
frequency of any given changes in individual plants.
Had it been possible to give statistics setting forth
the frequency of certain deviations in plants or groups
having a particular organisation, as compared with the
rarity of their occurrence in other plants of a different

488 COMPENSATION.

conformation, these co-relationships would have been
rendered much more evident. A hundred different
plants, for instance, may be named in any particular list,
of which fifty shall be of one type of structure, and the
remainder of another. And the co-relative changes
in each fifty may appear to be evenly balanced, but
so far is this from beimg 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 im the plants to whose names
it is affixed.

Compensation—But little further need be said on this
head. An atrophied condition of one part is generally
associated with an hypertrophied condition of another,
and scarcely a change takes place in one direction, but
it is associated with an inverse alteration in some other.
This principle is not universal, and its application must
not be unduly strained. It requires specially to be
considered in reference to differences in the degree or
kind of functional activity exercised by the organs
implicated--points beyond the scope of the present
volume.

Teratology and classification Lastly, there remain to
be mentioned. the bearings of teratology on systematic.
botany. There are those who would entirely exclude
teratology from such matters. It may be 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.

CLASSIFICATION. 489

Take the characters of exogens as distinct from
endogens ; even under ordinary circumstances, no abso-
lute distinction can be drawn betweenthem. There are
plants normally of an intermediate character, while, to
take exceptional instances, there are exogens with the
leaves and flowers of endogens, and endogens whose out-
ward organisation, at any rate, assimilates them to exo-
gens. Diclinous or monochlamydeous plants owe their
imperfect conformation to suppression, and may become
structurally complete by a species of peloria. Struc-
turally hermaphrodite flowers become unisexual by
suppression, or are rendered incomplete by the non-
development of one or more of their floral whorls.
Hypogynous flowers become perigynous by adhesion,
or by lack of separation; perigynous ones become
hypogynous by an early detachment from the 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 may be altered has already been alluded to, as has
also the circumstance that while it 1s common to find
an habitually imferior or adherent ovary becoming
superior or free, it is much more rare to find the
superior ovary adherent to the receptacle or to the
ealyx.' Regular and irregular peloria, too, serve
to show how slight are the boundaries, not only
between different genera, but also between different
families.

While, therefore, teratology may be an unsafe guide
in strictly artificial schemes, it 1s 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, yhich
was surmounted by the calyx lobes, precisely as in the case of a ; tha.
ceous fruit, has been given at p. 424, adnot.

490 CLASSIFICATION.

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 defined differentiation, we may assume
them to be evidences of a progressive rather than of a
retrogressive tendency.

That monstrosities so called may become the starting
points of new forms is proved by 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.

APPENDIX.

DOUBLE FLOWERS!

In ordinary language, the epithet double flowers is applied to flowers
of very varied structural conformation. The most common conditions
rendering a flower double, in the popular acceptation of the term, are
substitutions of petals or petal-like bodies for stamens and pistils, one
or both. (See Petalody, p. 283.) Another very common mode of
doubling is brought about by a real or apparent augmentation in the
number of petals, as by multiplication, fission, or chorisis. (See pp.
66, 343, 371, 376.) Sometimes even the receptacle of the flower within
the outer corolla divides, each subdivision becoming the centre of a new
series of petals, as in some very luxuriant camellias and anemones.
The isolation of organs which, under ordinary circumstances, are united
together, is another circumstance, giving rise, in popular parlance, to
the use of the term double flower. (See Adesmy, Solution, pp. 58,
76, 82.) Prolification is another very frequent occurrence in the case
of these flowers, while still other forms arise from laciniation of the
petals, or from the formation of excrescences from the petals or stamens,
in the form of supplementary petal-like lobes. (See Hnation, p. 443.)

As these matters are all treated of under their respective headings,
it is not necessary to allude to them again in detail. It may be well,
however, to allude, in general terms, to the causes which have been
assigned by various writers for their formation, and to the means which
have been adopted by practical experimenters to secure the production
of the flowers often so much esteemed by the florist. It must be
admitted that, in spite of all that has been written on the subject,
but very little is known about these matters. In the case of the stock
the following means have been adopted by cultivators in order to

1 This appendix forms a portion of a paper published in the ‘ Pro-
ceedings of the International Botanical Congress,’ London, 1866, p. 127,
and which it has been deemed advisable to reproduce with sundry
additions and modifications. :

492 APPENDIX.

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 im the plan of
planting them during the last quarter of the moon, but such confidence
is found to be misplaced. The plan of removing the stamens has had
its supporters, but as this must be done at an early stage of development,
and could only influence the result by diverting the vital force which
would be expended in the maturation of the pollen, to the perfecting of
the seeds, it is obvious that the plan is impracticable for all ordinary
purposes, even if in any degree efficient, which from the plasticity of
vegetable development, and the faculty of doubling which is inherent
in the stock family, is not at all improbable. Still another mark, the
presence of a fifth petal in the single or seed-bearing flower, has been
held to indicate the assurance of obtaining a crop of double-flowered
plants from seeds saved from flowers possessing this peculiarity. Toa
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. Chaté, who has written a
book! 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 ees furnish double flowers from those which would
produce ae ones.”

: ‘Teaité des ence, per E. Giare:

APPENDIX. 493

M. Chaté’s method, which he calls the French one, gives still greater
results, viz.: 80 per cent. of double flowers, and these produced by very
simple means. “When my seeds,” he observes, “have been chosen
with care, I plant them, in the month of April, in good dry mould, in a
position exposed to the morning sun, this position being the most
favourable. At the time of flowering I nip off some of the flowering
branches, and leave only ten or twelve pods on the secondary branches,
taking care to remove all the small weak branches which shoot at
this time. I leave none but the principal and the secondary branches
to bear the pods. All the sap is employed in nourishing the seeds
thus borne, which give a result of 80 per cent. of double flowers. The
pods under this management are thicker, and their maturation is more
perfect. At the time of extracting the seeds the upper portion of the
pod is separated and placed aside, because it has been ascertained that
the plants coming from the seeds situated in this portion of the pod,
give 80 per cent. of single flowers. They yield, however, greater
variety than the others. This plan of suppressing that part of the
pod which yields single flowers in the largest proportion, greatly
facilitates the recognition of the single-flowered plants, because there
remains to be eliminated from among the seedlings only from 10 to 15
per cent.

This separation of the single from the double-flowered plants, M.
Chaté 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.
Chaté’s method of securing so large a proportion of double-flowered
plants, and then of separating them from the remaining single ones—
a method which commends itself to the good sense of the intelligent
cultivator.”'

Signor Rigamonti, a great cultivator of pinks, asserted that he was
able to distinguish double from single-flowered pinks, in the seedling
state. According to this gentleman, those seedlings which produce
three cotyledons in a whorl in place of two, form double flowers.
In the case of Primula sinensis the same results occurred. Some had
three leaves in a ring, others two; most had the leaves standing one
over the other as usual. These were divided into three sets, and when
they flowered, the first lot were all double, the second semi-double, the
third single. But these statements have not been confirmed by other
observers; and the writer can safely assert that seedling pinks occa-

‘ Leading Article in the ‘Gardeners’ Chronicle,’ p. 74, 1866.

494. APPENDIX.

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-
flowered.

“Tn the year 1820 we several times visited a garden in the neighbour-
hood of Vienna, well known on account of its plant culture. The
gardener there possessed an immense plant of Caimellia japonica with
single flowers, and some small plants raised from this by cuttings, but
no other variety of camellia. He fertilised the flowers with their own
pollen, harvested seeds, which he sowed, and the plants raised from them
were placed in an extremely dry, lofty conservatory, where, after some
years, instead of producing single flowers, they all produced double
' ones. The seedlings and mother plant were planted in one and the same
kind of earth, and some of the flowers on the old plant also showed an
inclination to become double.

“This, at that time, to us, enigmatical phenomenon, was kept in mind
until we had an opportunity of instituting comparisons between the
climate of Japan and China and our own, and we then concluded that
in the case of a plant imported from thence, and exposed to such
different climatical influences, the origin of the greater or less 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

- APPENDIX. : 495

often transformed into deserts. Is it, therefore, remarkable that a plant
introduced from Japan into Europe, exposed to the influences of this
great diversity of climate, should produce imperfect sexual organs
incapable of further propagating the plant from seeds? A rich soil, with
the necessary amount of moisture, will never engender double flowers.”

Mr. Darwin? describes a peculiar form of Gentiana Amarella, in which
the parts of the flower were more or less replaced by compact aggrega-
tions of purple scales in great numbers. A similar condition is, indeed,
not uncommon in this plant, and, as Mr. Darwin also remarked, on
hard, dry, bare, chalky banks, thus bearing out the views expressed by
the writer in the ‘Gartenzeitung’ just cited. Some double flowers of
Potentilla reptans found growing wild near York, and transmitted to the
writer by a correspondent, were observed growing along a high wall,
in a dry border, close to a beaten path, bordering on a gravel pit,
others were found on a raised bank, which, from its elevation and expo-
sure to the sun, was particularly dry.

On the other hand, the double-flowered Cardamine pratensis, which is
occasionally found in a wild state, always grows in very wet places.

Of late years a remarkable double-flowered race of Primula sinensis
has been obtained. In particular, Messrs. Windebank and Kingsbury,
of Southampton, have succeeded in raising a set of plants in which the
flowers are very double and very attractive in a florist’s point of view.
The corollas in these flowers are not merely duplicated, but from their
inner surface spring, in some cases, funnel-shaped or tubular petals
(p. 315), so regular in form as quite to resemblea perfect corolla. These
tubes are attached to the inner side of the tube of the corolla, in the
same way as are the stamens, these latter organs being, it appears,
absent. The carpels are present, but open at the top, and bear nume-
rous ovules, hence it was at first surmised that these plants were
obtained and perpetuated, by the application of pollen from single
flowers to these double-flowered varieties.

The raisers of this fine race however assert that “the double kinds
are all raised from the seed obtained from single flowers; the double
blooms do not produce seed, as a rule, and even if they did yield seed,
and it were to germinate, the plants so raised would simply produce
single flowers.” Semi-double flowers will produce seed, but it is neces-
sary 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 Otto’s ‘ Gartenzeitung,’ 1866.
2 «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.!

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 go 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-fiowered 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.’
On these points, the evidence furnished by a double white hawthorn
in the Royal Botanie Gardens at Edinburgh is important. Professor

1 «Gard. Chron.,’ 1867, 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.”

* See De Candolle, ‘Plant. Rar. Genev.,’ 1829, p. 91; ana Alph. de
Candolle, ‘ Géog. Bot.,’ p. 1080.

APPENDIX. 4.97

Balfour kindly wrote as follows in reply to an inquiry respecting this
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 flowering has been always weak, since this change
of flowers from double to single. Mr. M‘Nab attributes the change in
the duration of the leaves to the filling up of the ground round the
tree, to the height of a foot and a half on the stem. He is now trying
the effect of extra manure in giving extra vigour to the plant.” Here,
at least, the production of single flowers would seem to be the result
of debilitating causes, connected with the unusual persistence of the
leaves, &c., for while the tree was healthy, double flowers were pro-
duced.

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 ijn 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 Ranwneulus 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-

32

4.98 APPENDIX.

jections raised by the present writer and others, and examines some
of the alleged exceptions. Some of these the Belgian savant finds to
prove his rule, inasmuch as although there is a co-existence of 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 cases in which double flowers
and variegated foliage do exist in conjunction, and where the plants
are vigorous and the flowers well developed. Instances of this are
known to cultivators in species of Dianthus, Hemerocallis, Althea,
Peonia, 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.’

Mr. Darwin? 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 ofa 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, E. 1115.
* Thid., 1843, p. 628.

APPENDIX. 4YY

tive action is stopped, at least partially—pretty much as it would be if
the plant were placed in the opposite condition of starvation. The
effect of supplying a plant (or an animal) with an excessive supply of
food, which it cannot assimilate,is in many respects similar to that
which results from partially cutting off the supplies. And the same
reasoning applies to sterility. If by high culture, or the supply of an
undue quantity of nourishment, the constitution of the plant be im-
paired, or if the plant be pampered, it is mo wonderful thing that sterility
shouldensue. 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 biade—are
present.

When once the disposition to form double flowers is established, that
tendency becomes hereditary ; there are races of single Stocks in which,
out of hundreds of plants, scarcely one double-flowered form is met
with ; but when the tendency to produce double blooms is set up, single
flowers become the exception: thus, in the Balsams, before mentioned,
not one in fifty now produces single flowers, and the seeds of these
double Balsams produce double-flowered seedlings, with scarcely a
“rogue” among them.

The following list of plants producing double flowers of any kind
is taken from that given in ‘ Seemann’s Journal of Botany,’ vol. ii, p. 177,
and to which some additions have been made. Miscalled double flowers,
such as those of the Composite, Viburnum Hydrangea, &c., are excluded.

RANUNCULACEZ.

Clematis Viticella, Linn., 8. Europe.
florida, Thunb., Japan.
Fortunei, Moore, Japan.
patens, Desne, Japan.

Anemone japonica, Sieb. et Zuce., Japan.
coronaria, Linn., S. Europe, Asia Minor.
hortensis, var. Linn., S. Hurope.
palmata, Linn., N. Africa, Spain, Portugal. ‘
nemorosa, Linn., Europe, N. America, Siberia.
sylvestris, Linn., S. Europe, Siberia.

OO APPENDIX.

Hepatica triloba, Chaiwz., Europe.

Ranunculus bulbosus, Linn, Europe, N. Amer.
repens, Linn., Europe, Siberia, N. Amer.
acris, Linn., Europe, Siberia.
aconitifolius, Linn., Europe.
gramineus, Linn., Italy, France, Portugal, Switzerland.
bullatus, Linn., S. Europe.
aslaticus, Linn., The Kast.

Ficaria ranunculoides, Mench., Europe. .

Thalictrum anemoides, Michx., N. America.

Caltha palustris, Linn., Europe, Asia, N. America.

Trollius europzus, 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.

Peonia Moutan, Sims, China, Japan.

officinalis, Retz., Europe.
tenuifolia, Linn., Tauria.
albiflora. Pall., Siberia.
paradoxa, Andr., S. Europe.

NYMPH ACER.

Nelumbium speciosum, Willd., Africa, Asia.

BERBERIDACE.
Berberis, sp. cult.

PAPAVERACES.

Papaver Rheeas, Linn., Europe.
bracteatum, Lindl., Russia.
somniferum, Linn., 8. Europe, Asia Minor, Egypt.
Chelidonium majus, Linn., Europe, Asia.
Sanguinaria canadensis, Linn., N. America.
Podophyllum peltatum, Linn., N. America.

CRUCIFERZ.

Mathiola incana, R. Br., Mediterranean.

glabrata, D. C.

annua, Sweet., South Europe, Syria.
Cheiranthus Cheiri, Linn., Europe.
Iberis umbellata, Linn., Europe.

amara, Linn., Hurope.

Cardamine pratensis, Linn., Europe, Asia, Africa, America.
Hesperis matronalis, Linn., Europe, Siberia.
Barbarea vulgaris, 2. Br., Europe.

APPENDIX. 5O1

Sinapis arvensis, Linn., Europe.
Brassica oleracea, Linn., Europe.

CISTACE®.

Helianthemum vulgare, Spach., Europe, N. Africa.

VIOLACEA,

Viola odorata, Linn., Europe, Siberia.
grandiflora, Linn., Europe.
tricolor, Linn , Europe.

CARYOPHYLLE.

Dianthus barbatus, Linn., France, Germany.
chinensis, D. C., China.
Poiretianus, Seringe, P
Caryophyllus, Linn., France, Italy.
arboreus, Linn., Crete.
hybridus (gardens),
corymbosus, Sibth., Asia Minor.
plumarius, Linn., Europe, Siberia, N. America.
deltoides, Innn., Europe.
Saponaria officinalis, Linn., Europe.
Lychnis sylvestris, Schkr., Europe.
vespertina, Linn., Europe.
flos cuculi, Linn., Europe.
Viscaria, Linn., Europe.
chalecedonica, Linn., Japan, Asia Minor.
Silene inflata, Sm.; var. maritima, D. C., Europe.

ALSINES.

Sagina procumbens, Linn,, Hurope.

MALVACE.

Hibiscus Rosa sinensis, Linn., H. Indies.
flavescens, Cav., China.
alba, Hook., China.
syriacus, Linn., Syria, Carniola.
Althea rosea, Cav., Caucasus, &c.
Malva rotundifolia, Linn., Europe.
moschata, D. C., Europe.

HIPPOCASTANES.

AXsculus Hippocastanum, Linn., Europe, N. America,

GERANIACE,

Geranium pratense, Linn., Europe, Siberia.
sylvaticum, Linn., Europe.
Pelargonium zonale, Willd., 8. Africa.
Tropzolum majus, Linn., Peru.
minus, Linn., Peru.

Ri

17
:

502 APPENDIX.

Oxalis cernua, Thumb., S. Africa,
Impatiens Balsamina, Linn., H. Ind.

TERNSTROMIACE®.

Camellia reticulata, Lindl., China.
Sasanqua, Thunb., China.
japonica, Linn., Japan.

Thea malifiora, Seem., Japan.

AURANTIACER,
Citrus Aurantium, Limm., Asia, South Europe.

PAPILIONACEZ.

Trifolium repens, Zinn., Europe, S. America.

Medicago sp., ., Europe.

Ulex europzus, Link., Europe.

Spartianthus junceus, Linn., S. Europe.

Clitoria Ternatea, Linn., E. India.

Orobus viscoides, D. C., Croatia, &e.
vernus, Linn., Europe.

Genista tinctoria, Linn., Burope.
sibirica, Iinn., Siberia.
scoparia, Lam., Europe.

Cytisus albus, Link., Portugal.

Anthyllis Vulneraria, Linn., Hurope.

Coronilla Emerus, D. C., Europe.

Lotus corniculatus, Linn., Europe.

ROSACEA.

Rosa lutea, Mill., Europe.
cinnamomea, Linn., Hurope, N. America.
spinosissima, Linn., Central Asia.
Carolina, Linn., N. America.
villosa, Linn., Europe, Central Asia.
centifolia, Linn.
damascena, Linn., Syria.
rubiginosa, Linn., Europe, Asia, N. America.
moschata, Ait., Madeira, N. Africa.
canina, Linn., Kurope.
alba, Linn., Europe, Caucasus.
indica, Linn., China.
nivea, D. C., China.
Eglanteria, Linn., Europe.
gallica, Linn., Europe, Caucasus.
pimpinellifolia, Linn., Europe, Central Asia,
Banksiz, R. Br., China.
sulphurea, Ait., East.
Rubus fruticosus, Linn., Europe.
rosifolius, Linn., Mauritius, E. India.
corylifolius, Smith, Europe.
cesius, Linn., Hurope.
Kerria japonica, D. C., Japan.

APPENDIX.

Spirea Filipendula, Linn., Europe.
Ulmaria, Iinn., Europe.
runifolia, Sieb. et Zucc., Japan.
eevesii, Lindl., China.
strobilacea, Sieb. et Zuce., Japan.
Fragaria vesca, Linn., Europe, N. America,
Potentilla alpestris, Hall. f., Europe.
reptans, Linn., Europe, Asia.
Tormentilla, Schrank, Europe, Asia.
anserina, Linn., Europe.
Geum rivale, Linn., Europe.

PoMACEZ,

Crategus Oxyacantha, Linn., Europe.
Crus galli, Linn., N. America.
Cydonia japonica, Pers., Japan.
Pyrus communis, Linn., Hurope.
Malus, Linn., Europe.
Eriobotrya japonica, Lindl., Japan.

AMYGDALER.

Amygdalus Persica, [inn., Persia.
communis, Jinn., Mauritania.
Prunus domestica, Linn., Europe.
spinosa, Linn., Europe, N. America.
avium, Linn., Europe.
Cerasus, Linn., Europe.
Kerii, Steud., Japan.
japonica, Thunb., China, Japan.
insititia, Linn., Hurope.
triloba, Lindl., China.

MYRTACES.
Myrtus communis, Linn., S. Hurope.
Punica Granatum, Linn., 8S. Europe, Marocco.
PHILADELPHACE.

Philadelphus coronarius, Linn., 8. Europe.
Deutzia crenata, Sieb. et Zuce., Japan.

ONAGRARIZ,

Fuchsia globosa, Lindl. (and vay. hort. pl.), Mexico.

Epilobium tetragonum, D,C., Europe.
Clarkia pulchella, Pursh., California,
elegans, Douglas, N, America.

PORTULACACES.
Portulaca grandiflora, Hook, Chili.

GROSSULARIACES.
Ribes sanguineum, Puish., N. America.

~
*

fe
ava

4)

503

504 APPENDIX.

SAXIFRAGACEA.

Saxifraga granulata, Linn., Europe.

UMBELLIFER 2.
Daucus Carota, Linn., Europe.

RUBIACEA.

Ixora grandiflora, De Cand., EB. India.

Serissa feetida, Comm., China, J apan.

Gardenia Fortuniana, Hook., China.
florida, Linn., China, E. India.
radicans, Thunb., Japan.

CAPRIFOLIACEA.

Lonicera Periclymenum, Linn., Europe.
Sambucus nigra, Linn., Europe.

CAMPANULACES.

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, [inn., Europe, Asia.
Medium, Linn., Europe.
rhomboidea, Linn., Europe.

Platycodon grandiflorum, D. C., Siberia.

ERICACEA.

Calluna vulgaris, Linn., Europe, N. America.
Rhododendron indicum, Sweet., BE. India.
ponticum, Linn., Asia Minor.
Azalea nudiflora, Linn., N. America.
glauca, Lam., N. America.
Arbutus Unedo, Linn., 8. Europe.
Erica Tetralix, Linn., Europe.
cinerea, Linn., Europe.
hyemalis, gardens.

KPACRIDACE.
Epacris impressa, R. Br., Australia.

PRIMULACE®.

Primula villosa, Jacq., Europe.
Auricula. Linn., Hurope.
denticulata, Smith, EK. India.
acaulis, Jacq., Europe.
elatior, Jacg., Europe.
prenitens, Ker. = sinensis, Lindl., China.

APPENDIX. 505

Lysimachia Nummularia, Roem et Schult., Europe.
Anagallis tenella, Linn., Europe.

JASMINACE®.

Jasminum officinale, Linn., 8. Europe.
Sambac., Ait., E. India.
hirsutum, Hook., China.
grandiflorum, Lindl., S. Europe.

OLEACE.

Syringa persica, Linn., Persia.
vulgaris, Linn., Europe, Persia.

APOCYNES.

Vinea minor, Linn., Europe.
major, Linn., Europe.
Nerium odorum, A7zt., E. India.
Oleander, Linn., S. Europe.
Tabernemontana coronaria, Willd., E. India.
Allamanda cathartica, Awbl., S. America.

CONVOLVULACES.

Calystegia sepium, R. Br., Europe, America, Asia.
pubescens, Lindl., China.

Convolvulus tricolor, Linn., S. Europe.

Ipomeea pandurata, Meyer, S. America.

SOLANACES.

Datura cornigera, Hook., Peru.
fastuosa, Linn., 8. America, Egypt.
arborea, Linn., S. America.
chlorantha, Hook.
humilis, Desf.
Petunia nyctaginiflora, Juss., S. America.
violacea, Hook, S. America.
Solanum Dulcamara, Linn., Europe.

GENTIANACE.

Gentiana Amarella, Linn., Europe.

OROBANCHACE®.
Orobanche sp.

ScROPHULARIACE.

Mimulus luteus, Linn., Chili.

Antirrhinum majus, Linn., 8. Europe.
Digitalis purpurea, Linn., Europe,

Linaria vulgaris, Mill., Europe, N. America.
Veronica, sp.

Calceolaria, var. cult.

506 APPENDIX.

GESNERACEZ.

Achimenes longifiora, D. C., Mexico.
Gloxinia var. hort.

VERBENACES.

Clerodendron fragrans, Willd., Japan.
Verbena var. hort.

NYCTAGINEZ.
Mirabilis Jalapa, Linn., Trop. America.

LAURINES.

Laurus nobilis, Linn., 8. Europe.
Sassafras, Linn., N. America.

IRIDACE.

Gladiolus tristis, Zinn., Cape of Good Hope.

Crocus aureus, Sibth, Europe, Asia Minor.
Susianus, Owrt., Asia Minor.
pusillus, Tenore, Italy.
vernus, Smith, 8. Europe.

Tris sibirica, Linn., Europe.

Tris Kempferi, Siebold, Japan.

AMARYLLIDACES.

Galanthus nivalis, Linn., Europe.
Leucoium vernum, Linn., Europe.
Sternbergia lutea, Gawl., Europe, Asia Minor.
Hippeastrum equestre, Herb., S. America.
Narcissus cernuus, Salisb., S. Europe.
Telamonius, Schult., Europe.
lobularis, Schult. x
concolor, Schult., Portugal.
biflorus, Curt., Europe.
italicus, Ker., Italy.
incomparabilis, Curt., Italy.
Cypri, Haw., Cyprus.
Pseudo-Narcissus, Linn., Europe.
poeticus, Linn., Europe.
Jonquilla, Linn., 5. Europe, Hast.
Tazetta, Linn., 5. Europe.
poculiformis, Salisb., S, Europe.

.

ORCHIDACES.

Orchis Morio, Linn., Europe.
mascula, Linn., Europe.
pyramidalis, Linn., Europe.

Ophrys fucifera, Linn., Europe.

See also pp. 380, 509.

HYDROCHARIDACE®.
Hydrocharis Morsus rane, Linn., Europe.

APPENDIX. 507

ASPHODELE®.
Asphodelus luteus, Linn., 8. Europe.

LILIACE.

Tulipa Gesneriana, Linn., Asia Minor.
sylvestris, Linn., 8. Europe.
Scilla autumnalis, Linn., Europe.
nutans, Smith, 8. Europe.
Convallaria majalis, Linn., Europe, America,
Polygonatum, Linn., Hurope.
Trillium grandiflorum, Spreng., America.
Fritillaria Meleagris, Iinn., Kurope.
imperialis, Linn., Persia.
Lilium Martagon, Linn., Hurope.
candidum, Linn., Syria, Persia.
Hyacinthus orientalis, Linn., East.
Polianthes tuberosa, Linn., E. India.
Hemerocallis disticha, Don., Nepal.
Kwanso, gardens.
fulya, Linn., 8. Europe.

CoLCHICACE.

Colchicum autumnale, Linn., Europe.
Tofieldia calyculata, Wahl., Kurope.

BUTOMACE®.
Sagittaria latifolia, Willd., N. America. '
sagittifolia, Linn., Europe, Asia, America.
COMMELYNACE.

Tradescantia virginica, Linn., N. America.
alba, gardens.

NOE.

Durinea the progress of the foregoing pages through the press,
several additional illustrations of particular malformations have come
under notice. Some of the more important of these may here be
recorded.

Fasciation (see p. 11).—The following plants may be added to the
list:—Acer eriocarpum, Arabis albida, Brassica oleracea, var., Guarea,
sp., Artabotrys sp. In all, with the exception of the first-named, the
fasciation occurred in the inflorescence. In some species of Artabotrys,
indeed, fasciation and curvation of the inflorescence are common.

Synanthy (p. 39)—Several additional instances of adhesion of two or
more flowers in Calanthe vestita, OC. Veitehti, 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 Crateegus
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
Crategus Oxyacantha present the same anomalies.” For analogous
instances in Digitalis, see p. 98. See also p. 580, Sawifraga.

Prolification, p. 120.—A. P. De Candolle, ‘“ Organographie Végétale,”
tab. 40, figures an instance of suppression of one lobe of the ovary in
Tris chinensis, and of the presence at the base of the flower of an

NOTE. 509

adventitious and imperfect flower-bud, as in the Phlomis, mentioned at
p. 119.

Monecious Misleto, p. 195.—In this specimen, exhibited at one of the
meetings of the Scientific Committee of the Royal Horticultural Society
in 1869, there were both male and female flowers on the same bush. The
plant was of the male sex, with numerous long slender whip-like, somewhat
pendulous, branches bearing comparatively large broad yellowish leaves,
and fully developed male flowers at the end. From the side of one of
these male branches, near the base, protruded a tuft of short, stiff-
branches, bearing small, narrow, dark green leaves, ripe berries and
immature female flowers. There was no evidence of grafting or 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. 380 and 355. In a species of
Picrasma, 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 pinne 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, 7. e., away from the corresponding portion of the neighbouring
pinne, 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
pinne, 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 labella in Phaius——A flower of Phaius grandiflorus was
found in the same condition as the Catasetwm, mentioned at pp. 291 and
382.

Tubular stem.—A species of Sempervivum, exhibited by Mr. Salter, of
Hammersmith, at one of the summer exhibitions of flowers at the Royal
Horticultural Society in 1868, under the name of S. Bollei, deserves
notice from its bearing on the question of such structures as the calyx-
tubes, the hip of the rose and such like, see pp. 394, 482. In this plant

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. 482.

Double flowers, see pp. 499, et seq.—The following species may be added
to those already recorded: Lychnis coronaria, Hibiscus mutabilis, Lotus
major, Pisum sativum, Godetia sp., Ipomea purpurea, Convolvulus minor,
Heliotropium peruvianum, Trillium grandiflorum, and Phaius grandi-

florus.

INDEX TO

A.

Abortion, 455, 467
of axile organs, 455
calyx, 460
corolla, 460
indusium, 467
leaves, 458
ovules, 466
perianth, 460
pistil, 464.
receptacle, 457
stamens, 463
Acaulescence, 393
Acaulosia, 393, 456
Acheilary, 398
Adesmy, 58, 76
Adhesion, 32
of axes, 50, 55
embryos, 56
leaves, 33
parts of flower, 34
roots, 53
Adveutitious buds, 156, 176
flowers, 174, 176, 180
gemme, 173
leaves, 162
roots, 156
shoots, 161
Albinism, 337
Alternation, 3, 485
Andrecium, enlargement of, 430
meiophylly of, 398
meiotaxy of, 405
of orchids, 380
pleiotaxy of, 379
polyphylly of, 361
Androgynism, 193
Anther (see connective), contabescence
of, 463
morphology of, 291
ovuliferous, 200
petalody of, 291
sutures of, 291

SUBJECTS.

| Apilary, 397

Apostasis, 440

Aphylly, 395

Arrangement, 1

Ascidia, 30, 313 .
bibliography, 30
plants with, 30

Atrophy, 454 (see abortion)

Autophyllogeny, 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

B.

Barley, Nepaul, 174:
Bigarades cornues, 303
Bladder-plums, 465
Bracts, multiplication of, 358
staminody of, 298
phyllody of, 24.2
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-
lification)
variations of, 336
Bulbs, displacement of, 84
multiplication of, 172, 350
Burrs, 347, 420

C.

Catacorolla, 450
Calycanthemy, 283

912 INDEX
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, 3438
parallel, 344
collateral, 344
Chromatism, 339
Cladodes, 328

Classification, teratology in relation to,

488
Cohesion of leaves, 21, 25
of petals, 28
pistils, 29
sepals, 27
stamens, 29
stems, 9
Colour, alterations of, 337
Columella, 395
Conifere, leaves of, 217, 362, 454:
inflorescence of, 245
scales of, 192, 245, 484
Compensation, 488
Connective, petalody of, 293
Consistence, alterations of, 432
Contabescence, 463
Contortion, 317
Co-relation, 486
Cornute leaves, 328
Corolla, abortion of, 461
dialysis of, 71
duplicate, 376
hose in hose, 377
meiophylly of, 397
meiotaxy of, 403
pleiotaxy of, 374
polyphylly of, 359
virescence of, 338

TO SUBJECTS.

Cotyledons, increased number of, 370
shoots, below, 161
Cuttings, formation of, 159

D.

Dédoublement, 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
Dicecious plants, 192, 193
Dimorphism, 333
Dimorphice flowers, 403
Direction, changes of, 201
Diremption, 87
Disjunction, see Dialysis, Fission, So-
lution
Displacement, 84
of bulbs, 84
carpels, 96
flower-parts of, 91
inflorescence, 85
leaves, 87
ovules, 96
placentas, 96
Distension, 419

E.

KEcblastesis, 107, 138
Elongation, 433
of parts of flower, 438
flower-stalks, 435
inflorescence, 434
leaves, 437
nucleus of ovule, 269
placenta, 440
receptacle, 440
root, 434
thalamus, 440
Embryos, adhesion of, 56
increased number of, 369
Enation, 443
from axile organs, 444.
carpels, 453
corolla, 449
foliar organs, 445
sepals, 448
stamens, 453

INDEX TO

Enlargement, 417
of andreecium, 430
axile organs, 418
buds, 420
flower stalk, 421
fruit, 431
gynecium, 430
perianth, 428
placenta, 425
Epanody, 226
Epistrophy, 226
Etiolation, 337
Eversion, 204
Excrescences, 444 (see Enatiou)

F.

Fasciation, 11
plants affected with, 20, 508
Fastigiation, 202
Ferns, crested, 63, 447
depauperated, 466
exindusiate, 467
supra-soriferous, 189
Filaments, see Stamens
petaloid, 290
4-winged, 290
Finger and toe, 69
Fission, 59
of carpels, 68
leaf-organs, 61, 66
plants, subject to, 66
of petals, 66
stem-organs, 60
stamens, 68
Flattening, 328
Floral organs, displacement of, 91
elongation of, 439
- metamorphy of, 281
Florets, increased number of, 351, 390
Flowers, adventitious, 174
on fruits, 177
on leaves, 174
in ovary, 180
on petals, 177
on spines, 177
apetalous, 404.
double, 490, 510
hermaphrodite, 196
homomorphic, 188
increased number of, 390
mutilated, 403
unisexual, 193
Flower-stalk, enlargement of, 421
Flower-bud (see Prolification), replaced
by leaves or scales, 164.
in place of leaf-buds, 176
Foliar organs (see leaves), adhesion of,
32

SUBJECTS.

513

Foliar organs, cohesion of, 21, 25
enation from, 445
fission of, 61
suppression of, 396
Form, alterations of, 213
juvenile, persistence of, 217
Frondescence, 241, 279, see Phyllody,
Virescence
Fruit, adhesion of, 44
enlargement of, 431

G.

Gemme, formation of, 173
Glands, formation of, 473
Gnaurs, 158, 347, 417, 419
Grafting, 53, 56
Greffe des Charlatans, 56
Growth interrupted, 327
irregular, 228
Gymnaxony, 211
Gynantherus, 305
Gynecium (see pistils)
enlargement of, 430
meiophylly of, 399
meiotaxy of, 406
pleiotaxy of, 388
polyphylly. of, 363
suppression of, 406

H-

Hairs, formation of, 472
Hermaphroditism, 197
Heterogamy, 190

Heteromorphy, 311

Heterophylly, 330

Heterotaxy, 156

Homology, 476

Homomorphy. 188

Hose in hose corollas, 291, 377
Hypertrophy, 415, see Enlargement

if

Independence, 58

Indusium, abortion of, 467

Inflorescence, displacement of, 84
elongation of, 434
prolification of, 102, 115

Interrupted growth, 327

Inversion of organs, 206

Irregularity, 213

“Irregular growth, 228

kK.

Knaurs, (see gnaurs)
Kail, 426
35

E

514

L.

Laciniation, see fission

Layering, 156

Leaders, formation of, 203

Leaf-sheath, 477

Leaves, see foliar organs
abortion of, 458
adhesion of,

by surfaces, 33
to stem, 34

adventitious, 162—165, 509
cornute, 328
displacement of, 86
elongation of, 437
enlargement of, 421
frondiferous, 355
geminate, 3852 —
multiplication of, 358
nature of, 477
palmate-passage of to pinnate, 439
spiral torsion of, 326
supernumerary, 3853

Lily, double white, 375

M.

Meiotaxy of andreecium, 405
of calyx, 403
corolla, 4.03
gyneecium, 405
Meiophylly of andrecium, 398
of calyx, 897
corolla, 397
gynecium, 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
Monecious, 192, 193, 509
Monosy, 58
Morphology, 479
Multiplication, see pleiotaxy,
phylly
of bracts, 358, 371
bulbs, 350
cotyledons, 370
embryos, 369
florets, 351
foliar organs, 352
whorls, 371

pleio-

INDEX TO

SUBJECTS.

ING

Nepaul Barley, 174
Number, alterations of, 341
increased, 343, 353

diminished, 392

O.

Orchids, andreecium 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

Ne

Parasitical plants, 55
Peduncles, elongation of, 4385
Peloria, 207, 228
bibliography of, 227, 239
regular, 219
plants subject to, 226
irregular, 229
plants, subject to, 239
Perianth, abortion of, 460
enlargement of, 428
meiophylly of, 396
pistillody of, 303
pleiotaxy of, 375
Persistence, xxxvi adnot., 217
Petalody, 283
of accessory organs, 297
anther, 291
calyx, 283
connective, 292
ovules, 297
pistils, 297
stamens, 284
plants, subject to, 295
Petals, cohesion of, 28
enation from, 448
fission of, 66
phyllody of, 251
staminody of, 298
tubular, 23, 314
Phyllode, 328

INDEX

TX )

Phyllody, 240 (see Virescence, Chlo-

ranthy,
of accessory organs, 272
bracts, 242
calyx, 244
plants subject to, 250
in Conifers, 245
of corolla, 251
plants subject to, 252
of ovules, 262
plants subject to, 271
pistils, 256 :
plants subject to, 261
stamens, 253
plants subject to, 256
Phyllomania, 352
Phyllomorphy, see Phyllody
Phyllotaxy, 1, 320
Pistil, abortion of, 464
cohesion of, 29
petalody of, 296
staminody of, 298
Pistillody, 302
of ovules, 268, 810
perianth, 302
sepals, 302
stamens, 303
plants subject to, 310
Pitchers, 30, 313, 509, see Ascidia
Placentation, changes in, 96, 508
bibliography, 100
nature of, 483
Placenta, elongation of, 439
enlargement of, 421
Pleiomorphy, 228
Pleiophylly, 353
Pleiotaxy, 371
of andrcecium, 375
bracts, 371
calyx, 374:
corolla, 376
gynecium, 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 andreecium, 361
of calyx, 359
corolla, 359
plants subject to, 360
of flower, 363
gyneecium, 363

|

|

|
|
|

| Saint Valery Apple, 135, 282, 304, 375,

SUBJECTS.

Polyphylly, bibliography, 364°
plants subject to, 364
Position, changes of, 83
relative, 485
Prolification, 100
axillary, 138
foliar, 141
floral, 142

ay
eo

plants affected with, 148

bibliography of, 154:

complicated, 151

of embryo, 155

of flower, 115

coincident changes, 128
median foliar, 116
median floral, 119, 508

plants affected with, 137

of fruit, 134
inflorescence, 102
bibliography, 115
median foliar, 103
median floral, 105
lateral floral, 107
lateral foliar, 106

R.

Receptacle, abortion of, 457
elongation of, 116, 440
spiral torsion of, 325

Reflexion, 209

Regularity, 213

Rhizotaxy, 1

Rose Willow, 166, 168

Roots, adventitious, 156
elongation of, 434

Rudimentary organs, 469

8.

388
Savoys, 426
Scales, formation of, 164, 448, 470
Scape, leaves on, 163
Seeds, abortion of, 407
union of, 50
Sepals, adhesion of, to petals, 34
cohesion of, 27
enation from, 448
phyllody of, 243
pistillody of, 303
staminody of, 298
Sepalody, 282
Separation, 58
Sex, changes of, 190, 509
Shamrock, four-leaved, 356
Shoots below cotyledons, 167;
Leaders

see

516

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 Andreecium
abortion of, 463
adhesion of, 34, 35
cohesion of, 29
compound, 294, 345
dialysis, 73
enation from, 453
fission of, 68
in ovary, 183
petalody of, 283
phyllody of, 253
pistillody of, 303
tubular, 316

Staminody, of accessory organs, 301

of bracts, 298
petals, 298
pistils, 299
sepals, 298
Stasimorphy, 216
Stem, see axes
Stipules, increased number of, 357
Strawberry, Plymouth, 275
blind, 195
Suppression, 393
of andreecium, 405
flower, 408
foliar organs, 395

INDEX TO SUBJECTS.

Suppression of ovules, 407
remarks on, 409
of seeds, 407

Symmetry, 213

Synanthy, 37
bibliography, 45

plants subject to, 44, 508

Synearpy, 45
Syngenesia, 29
Synophty, 57
Synspermy, 50

E.

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, Solenaidy, Spurs)

Tubular petals, 314
stamens, 316

U.

Union, 8
Unisexuality, 195
Uovoli, 420

We

Varieties, dwarf, 411
Venation, 338
Virescence, 338

Viviparous plants, 106, 168

Wie
Warts, 444

’ Wheat-ear carnation, 371

INDEX OF NAMES OF PLANTS.

[In the following Index the names of the orders that are 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, &e., 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 ata 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, &e., as
contrasted with other closely allied, and perhaps equally common species, under
apparently identical conditions, is also made manifest. ]

A. Adoxa, 367
Aerides odoratum, 398
Abies Brunoniana, 245 Aschynanthus, 44, 297
excelsa, 21, 61, 192, 298, 325, 456 | Adsculus Hippocastanum, 50, 66, 295,

Larix, 90 369, 438, 459, 501
pectinata, 52 Affonsea, 364
Acacia, 329 Afzelia, 397

Aceras anthropophora, 238
Acer, 359, 364, 367, 508
platanoides, 66, 459

pseudo-platanus, 20, 371 -

Aceranthus, 225, 226
Achimenes, 106, 114, 296
longiflora, 506

Aconitum, 148, 231, 359, 360, 399, 404,

407, 464
Napellus, 39, 44, 288
Acorus, 225
Actea spicata, 251, 252
Adenium obesum, 417

Adenorophium luxurians, 254

Adonis, 262, 295
autumnalis, 500
vernalis, 500

Agaricus, xxiii, 54
Agave, 177, 432
Americana, 33, 45
Angelica, 365
Agrimonia, 137, 406
Agrostemma, 148
Githago, 271
Agrostis alba, 169
AiJanthus glandulosa, 21
Aira alpina, 169
czspitosa, 169
Ajuga Iva, 404
pyramidal, 20
reptans, 243
Alcea, 149, 297
Alchemilla minima, 171

| Aldrovanda vesiculosa, 86

518 INDEX OF NAMES OF PLANTS.

Alisma, 115, 329
natans, 107
parnassifolia, 167
ALISMACE®, 115
Allamanda, 296
cathartica, 288, 390, 505
Allium, 106, 114, 170, 299, 360, 365,
367
fragrans, 369
vineale, 150
Alliaria officinalis, 269, 271
Almond, see Amygdalus.
Alnus, 349
fruticosa, 192
imperialis, 459
incana, 21
glutinosa, 66, 244, 349
laciniata, 65
Alopecurus pratensis, 169
Alsine media, 67, 252, 404.
Alstrémeria, 319, 326
Althea, 295
rosea, 20, 501
Alyssum, 137
Incanum, 252
AMARYLLIDACE®, 115, 138, 150
Amaryllis, 307, 310, 432
Ambrina ambrosioides, 397
AMENTACE®, 114, 435
Amorpha, 21, 397
fruticosa, 23, 30, 243
AMYGDALE®, 500; see Rosacea
Amygdalus, 122, 187, 155, 295, 297
364
communis, 250, 252, 369, 503
Persica, 74, 176, 503
Anagallis, 73, 138, 141, 150, 288, 296,
360, 365, 367
arvensis, 117, 161, 256, 263, 271,
278, 281, 284,
collina, 44
pheenicea, 141, 258, 271, 441, 461
tenella, 505
Webbiana, 281
Anomodon alternatus, 49
Ananassa, 350
Anchusa, 132, 138, 339, 365
ochroleuca, 125, 259, 262, 281
paniculata, 262
Androsace maxima, 16, 20
Anemiopsis californica, 63, 66
Anemone, 113, 121, 186, 140, 142, 148,
208, 289, 295, 296, 297, 360,
374, 430, 463, 491 (see Hepatica)
coronaria, 107, 250, 256, 339, 499
hortensis, 107, 250, 339, 499
japonica, 161, 499
nemorosa, 250, 256, 339, 499
palmata, 499

4

| Anemone payonina, 499

Pulsatilla, 250
rivularis, 165
sylvestris, 250, 499
Angelica, 114, 137, 148, 150
Razoulzii, 244, 437
Anthemis arvensis, 20
nobilis, 20
retusa, 44
Anthoxanthum, 61
odoratum, 61
Anthriscus, 442
Anthurium Scherzerianum, 358
Anthyllis, 295
Antirrhinum majus, 20, 22, 24, 31, 44,
57, 73, 82, 104, 114, 121, 131,
225, 226, 227, 230, 233, 235,
238, 253, 272, 296, 301, 315,
316, 318, 363, 365, 378, 398,
505
Apargia, 114,
autumnalis, 20
Apium, 82, 113, 150
graveolens, 66, 158
Petroselinum, 370, 437.
APOCYNACER, 137
Apple, St. Valery, 135, 282, 304, 375,
388

Aquilegia canadensis, 500
Skinneri, 266, 271
vulgaris, 24, 70, 127, 136, 220,
226, 252, 257, 260, 261, 271,
280, 286, 287, 288, 298, 295,
374, 390, 500
Arabis, 148, 508
alpina, 397, 461, 463
pumila, 170
sagittata, 44,
Araucaria, 245
Arbutus Unedo, 291, 292, 296, 377,
504:
Archidendron, 365
Ardisia serrulata, 369

| Arenaria serpyllifolia, 461

tetraquetra, 398, 399, 464, 469
Armeria, 114
Aristolochia Clematitis, 38, 45, 314.
caudata, 231, 23
sipho, 23
Armoracia rusticana, 64, 299
Arnoseris, 114
Artabotrys, 456, 508
Artemisia, 405
Artocarpus, 407
Arum, 329
maculatum, 66, 225, 227, 24.5, 358
Asparagus officinalis, 12, 19, 21, 320,
325, 471
Asphodelus, 138, 296

INDEX OF

Asphodelus luteus, 506
ramosus, 298, 310
Asplenium 'Trichomanes, 190

Astrantia, 114

major, 368
Athamanta, 82, 1387, 149

Cervaria, 250, 442
Atriplex, 227

hortensis, 224
Atropa Belladonna, 44
Aucuba, 21

japonica, 21, 197
AURANTIACER, 137, 149, 502
Avena, 319, 351, 391

chinensis, 298
Azalea, 35, 114

glauca, 504

NAMES OF PLANTS. 019

indica, 44, 73, 166, 209, 289, 296 |

nudiflora, 504

B.

Babingtonia, 185
Beckea diosmifolia, 183
BAaLsaMINED, 359, 404, 501
Balsam, see Impatiens
Bamboo, see Bambusa
Bambusa, 307, 310, 324, 365
Barbarea, 295

vulgaris, 310, 500
Barkhausia, taraxacifolia, 20
Barley, Nepaul, 174
Bauhinia, 328
BEGONIACER, 114

Begonia, 31, 81, 106, 114, 162, 170,

352
frigida, 199, 303, 310
fuchsicides, 281
phyllomaniaca, 170
Bellevalia, 408, 461, 467
comosa, 348

Bellis perennis, 17, 20, 31, 106, 114

164, 244.

Berberis, 272, 295, 458, 460, 500

vulgaris, 20
Beta, 19, 325, 365
vulgaris 20
Betonica Alopecuros, 42, 44, 226
Betula alba, 66, 346, 456, 472
populifolia, 66
Bidens, 114, 165, 223
BIGNONIACER, 222
Bignonia, 272, 296, 327
Bikkhia, 80
Blitum, 45
polymorphum, 397, 458
Bocconia, 224
cordata, 310

Bowiea volubilis, 409
BORAGINACER, 132, 138
Bouchea hyderabadensis, 166
Bougainvillea, 339
Brachythecium plumosum, 49
Brassica, 20, 1386, 139, 295, 364, 367
Napus, 27, 205
oleracea, 30, 33, 66, 67, 106, 113,
136, 141, 142, 148, 250, 252,
264, 271, 280, 351, 408, 421,
4.26, 442, 445, 501, 508
Rapa, 181
Breynia, 198
Bromelia, 103
Bromus velutinus, 358
Broussonettia papyrifera, 331, 459
Bruniacez, 80, 81
Brunia microphylla, 81
Bryonia, 360, 367
Bryophyllum calycinum, 158, 171, 483
proliferum, 166
Bryum czespititium, 49
Bunias, 97, 136, 280, 360
Bunium creticum, 159
flexuosum, 20
Bupleurum, 149, 459
faleatum, 15, 20
BUTOMACEX, 507
Buxbaumia indusiata, 49
Byrsonima, 137
BYTTNERIACE®, 362

C.

Cabomba aquatica, 458
CacrTacEes#, 81, 113, 149, 395
Cactus, 160
Cachrys taurica, 197
Ceesalpinia, 365
digyna, 48
Cakile maritima, 246, 250
Calanthe, 227, 398, 402, 508
vestita, 39, 45, 227, 402, 508
Caleeolaria, 41, 44, 230, 233, 284, 296,
397, 405, 406, 505
crenatifolia, 238
floribunda, 316
rugosa, 238

- Calendula, 114, 138, 339, 370

officinalis, 252, 280, 339
Calla palustris, 357
Callitriche, 196

antumnalis, 406

vernalis, 406
Caltha, 136, 148, 295

palustris, 141, 250, 442, 500
Calluna, 296, 504
Calycophyllum, 249, 283, 429
Calystegia, 114, 296

520 INDEX OF

Calystegia Sepium, 505
pubescens, 505
Camellia japonica, 288, 295, 297, 491,
494, 502
reticulata, 502
Sasanqua, 502
CAMPANULACER, 80, 114, 127, 131,
138, 139, 150, 334
Campanula, 71, 72, 73, 82, 138, 150,
250, 285, 296, 365, 367, 404,
442, 472
canescens, 403
colorata, 403
glomerata, 242, 300, 504
latifolia, 504
Medium, 20, 37, 44, 61, 251, 448,
504
persicifolia, 44, 284, 300, 429, 504
pyramidalis, 281, 504
Rapunceulus, 429
rapunculoides, 20, 252, 300, 310,
375
rhomboidea, 504
rotundifolia, 377, 378, 504:
Tenorei, 504
thyrsoidea, 20
Trachelium, 504
Vidallii, 504
Campanumea, 80, 81
Camphorosma monspeliaca, 456
Canna, 285
Cannabis, 82, 194, 197
sativa, 81
Cannamois virgata, 197
CAPPARIDACE®, 148, 390
CAPRIFOLIACER, 45
Capsella bursa pastoris, 298, 361
Cardamine, 295, 357
hirsuta, 398
Impatiens, 404.
pratensis, 65, 170, 181, 495, 500
sylvatica, 398
Carduus arvensis, 20
crispus, 166, 339
heterophyllus, 260, 262, 250
tataricus, 250, 260, 262
Carex, 115, 138, 150, 191, 194, 350
acuta, 143, 198
ceespitosa, 199
glauca, 143, 199
maritima, 369
paludosa, 199
vulpina, 428
Carica Papaya, 199
Carlemannia, 398
Carlina, 114
vulgaris, 20
Carpinus, 346
Betulus, 66

NAMES OF PLANTS.

Carthamus, 1388
Carum, 82, 114, 365
Bulbocastanum, 159
Carui, 244, 271, 285, 339, 437
CARYOPHYLLACER, 99, 113, 120, 137,
139, 140, 148, 250, 379, 397,
398, 404, 406, 407, 410, 418,
442, 443, 448
Casuarina rigida, 325

| Cassia, 364, 369

marylandica, 30
Castanea vesca, 11, 66, 104, 114, 319,
435
Catabrosa aquatica, 351
Catalpa, 399
Catasetum, 291, 296, 334
eburneum, 384
Cattleya amethystina, 401
Forbesii, 34, 384, 398
marginata, 223, 22
Mossiz, 224, 227, 238
violacea, 383, 397
Canealis leptophylla, 33
Caulophyllum. 75, 125
Caylussa, 137
Cedrus Libani, 61
CELASTRACEM, 149
Celastrus, 149
Celosia, 19, 20
Centaurea, 37, 114
collina, 34
Jacea, 43, 243
Scabiosa, 20
moschata, 44
Centranthus, 247
macrosiphon, 250
ruber, 42, 4:4
Cephalotus follicularis. 314
Cerastium, 62, 97, 262, 397, 398
glomeratum, 280, 358, 463
tetandrum, 4.63
triviale, 262, 280
vulgatum, 252, 404.
Cerasus, 74, 117, 149, 250, 260, 424,
489
avium, 262
caproniana, 364.
Lauro-cerasus, 64, 66, 370
vulgaris, 252, 262
Ceratonia Siliqua, 30
Cercis, 364 2
siliquastrum, 325
Chamerops humilis, 300
Cheerophyllum, 113

Cheiranthus, 121, 131, 136, 148, 295

364
Cheiri, 20, 33, 35, 36, 250, 252,
271, 404, 427, 500
var. gynantherus, 305, 310

a

INDEX OF

Cheiranthus incanus, 250
Chelone, 361
barbata, 238
Chelidonium majus, 66, 170, 280, 295,
500
CHENOPODIACE®, 397, 405, 406
Chenopodium, 45, 365, 367
glaucum, 397
murale, 428
Quinoa, 62, 66
Vulvaria, 458
Chirita sinensis, 170
Chlorophytum Sternbergianum, 107
Chorozema ilicifolium, 21
Chrysanthemum, 16, 72, 365
indicum, 20, 188, 472
Leucanthemum, 20
Chrysosplenium, 367
Cichorium Intybus, 20, 44
Cicuta virosa, 406
CINCHONACE, 429
Cionidium Moorei, 190
Cirsium, 114, 138
arvense, 250, 457
lanceolatum, 20
tricephalodes, 252, 339
Cissus, 211
CISTACE®, 137
Cistus vaginatus, 473
Citrus, 137, 149, 364, 453
Aurantium, 33, 35, 44, 56, 75,
134, 303, 310, 335, 369, 388,
389, 391, 502
Clarkia, 24, 295
elegans, 177, 503
pulchella, 503
Cleistanthus polystachyus, 198
Clematis, 136, 148, 288, 295, 367
florida, 499
Fortunei, 499
patens, 499
Viticella, 28, 499
Cleome, 137, 148
Cleonia lusitanica, 238
Clerodendron fragrans, 506
Cliffortia, 396
Clinacium dendroides, 49
Clitorea Ternatea, 231, 238, 295, 502
Clusia rosea, 11
Cluytia semperflorens, 198
Cneorum, 365
Cnicus palustris, 20
Cnidium, 113
Cobza scandens, 73, 82, 272, 326, 365
Coccoloba platycladon, 328
Cochlearia Armoracia, 64, 299, 310, 331
Cocos, 115, 365
nucifera, 429

NAMES OF

52]

Codixum variegatum, 31, 314, 326,
328, 459
Coelebogyne ilicifolia, 369
Cola acuminata, 370
Colchicum autumnale, 45, 67, 73, 250,
287, 296, 406, 407, 507
Coleus, 365, 459
Columnea Schiedeana, 226
Columbine, see Aquilegia
Colutea, 465
Commelyna, 73, 296, 507
COMMELYNACE®, 245, 507
Composite, 72, 73, 86, 107, 114, 127,
131, 188, 165, 223, 226, 235,
339, 406, 407, 430, 434, 437,
439, 442
Conceveiba macrophylla, 198
CoNIFER®, 56, 65, 103, 114, 191, 245,
369, 435
Conium maculatum, 114
Conostephium, 120
Convallaria maialis, 73, 150, 250, 296,
360, 367, 375, 400, 442, 507,
Polygonatum, 507
CONVOLVULACE®, 114, 137, 150
Convolvulus, 73, 114, 137, 142, 150,
296, 510
arvensis, 20
Sepium, 20, 108, 250
tricolor, 505
Conyza squarrosa, 20
Corallorhiza innata, 288
Circeia, 410
Corchorus acutangulus, 397
Coreopsis, 20, 73, 114, 138
Drummondi, 339
Cornus, 37, 44, 358
mas, 358, 374,
sanguinea, 44:
suecica, 374
Coronilla, 106, 118, 149, 295
Emerus, 502
Correa, 72, '73, 370
Cortusa Mathioli, 133, 138, 263
Corydalis aurea, 280
solida, 243
tuberosa, 235, 336, 238
Corylus Avellana, 21, 31, 48, 66, 114,
349, 354, 364, 365, 368
Cotoneaster microphylla, 21
Cotula feetida, 19, 20
CRASSULACE&, 113
Crassula, 113
arborescens, 26, 31
Crategus, 82, 113, 149, 295, 364, 404,
419
Crus galli, 503
monogyna, 44, 400

PLANTS.

522

INDEX OF NAMES OF

Crategus Oxyacantha, 57, 66, 78, 317, |

370, 503, 508
tanacetifolia, 107, 162
Crepis, 271
virens, 20
Crinum, 432
Crocus, 29, 35, 45, 67, 287, 289, 296,
319, 361, 399, 400, 434, 462
aureus, 006
nudiflorus, 302
pusillus, 506
vernus, 506
Crozophora tinctoria, 198
CRUCIFERH, 73, 76, 98, 113, 136, 139,
141, 148, 257, 297, 364, 379,
406, 410, 428, 442, 500
Cryptomeria japonica, 103, 114, 245,
435
Cucubalus, 149
Cucumis, 36, 82, 138, 248, 259, 326,
367
CUCURBITACER, 71, 80, 81, 187, 247

|
j

|
j
|
j
|
|
j

|

Cucurbita, 197, 201, 250, 307, 310, 365,

474 F
Cuphea miniata, 211, 424:
Cupressus funebris, 218
CrcaDE®, 56, 369
Cyclamen, 67, 104, 114, 296, 319, 359,
360
linearifolium, 329
Cyclodon, 80
Cydonia vulgaris, 71,79, 295, 423
japonica, 503
Cynanchum fuscatum, 369
nigrum, 369
Cynosurus cristatus, 169
CYPERACE®, 115, 138, 150, 169, 350
Cypripedium, 27, 92, 381, 386
candidum, 401
Hookere, 386
insigne, 91
Cyrtanthus, 177
Cytisus, 113, 295, 336
albus, 502
Laburnum, 21, 66, 157, 189, 222,
226, 231, 238, 356, 459
nigricans, 15, 104

D.

Dactylis, 115
clomerata, 169

Dahlia, 44, 72, 188, 244, 433
variabilis, 20

Danae, 60

Daphne indica, 21
odora, 21

Datura, 285, 291, 296, 363, 375, 407

PLANTS.

Datura arborea, 505
cornigera, 505
chlorantha, 505
fastuosa, 296, 379, 450, 505
humilis, 505
Daucus Carota, 53, 57, 82, 113, 12],
124, 125, 1387, 149, 244, 250,
252, 256, 260, 262, 296, 298,
339, 365, 368, 369, 370, 457,
504:
Delphinium, 44, 136, 148, 225, 283,
286, 295, 344, 364, 367, 374,
388, 399, 407
Ajacis, 250, 252, 261, 271, 339, 500
amcenum, 261
cheilanthum, 500
Consolida, 373, 500
crassicaule, 252, 256, 261, 271, 339
dictyocarpum, 271, 432
elatum, 20, 126, 237, 238, 261,
267, 271, 309, 310, 339
elegans, 500
grandiflorum, 500
peregrinum, 219, 226
Dendrobium, 227
nobile, 94, 398
normale, 224, 383
Desmodium canadense, 271
marylandicum, 467
Deutzia, 295
crenata, 503
Dianthus, 67, 113, 121, 129, 137, 139,
145, 146,.149, 166, 261, 268,
289, 295, 297, 310, 360, 364,
371, 379, 397, 471
arboreus, 501
barbatus, 325, 404, 501
Caryophyllus, 501
corymbosas, 501
deltoides, 501
hybridus, 501
plumarius, 501
Poiretianus, 501
sinensis, 370, 501
Dictamnus, 121, 122, 137, 140
albus, 256, 271
Fraxinella, 252, 262, 271, 278, 280
Dielytra, 286, 237
Digitalis lutea, 60
orientalis, 238
purpurea, 20, 40, 44, 73, 98, 12],
129, 137, 150, 226, 283, 238,
296, 298, 315, 365, 373, 398,
459, 472, 505
DILLENIACER, 398
Dionza, 308
Muscipula, 310
Diosma, 369
Diphaca, 365

INDEX OF

Diplotaxis, 136, 148, 364
muralis, 252, 458
tenuifolia, 73, 250, 261, 274, 280,
397, 398, 430
Dripsacace®, 86, 107, 114, 188
Dipsacus, 419, 429

NAMES

|
|
|

fullonum, 20, 62, 66, 281,*320, |

821, 325
Gmelini, 325
pilosus, 20, 325
sylvestris, 10, 20
Dipterocarpus, 249
Ditaxis lancifolia, 380
Dodecatheon, 138
Dodonza viscosa, 20
Draba, 364
Dracocephalum austriacum, 238
moldavicum, 20
speciosum, 320, 325
Dracontium pertusum, 25
Drosera intermedia, 170, 265, 271, 277,
473

E.

Ebenus cretica, 26
Eccremocarpus scaber,
Echeveria, 113
Echinophora maritima, 252
Echium pyrenaicum, 20

simplex, 20

vulgare, 374
Echinocactus, 149, 178, 417
Elegia, 115
Empetrum nigrum, 197
Encamptodon perichetialis, 174
EPACRIDACE®, 120, 137
Epacris, 103, 137

impressa, 61, 379, 504
Epidendrum, 114

elongatum, 107

Stamfordianum, 401
Epimedium, 23, 226

Musschianum, 390
Epiphyllum, 328
Epipactis palustris, 325
Epilobium, 81, 137, 273

angustifolium, 20

hirsutum, 246, 250, 252, 256, 262,

281, 442

palustre, 271

tetragonum, 503
Episcia bicolor, 170
EQUISETACE®, 189, 350
Equisetum, 61, 325

fluviatile, 325

limosum, 325

Telmateia, 323, 325

999,

mt

226, 326

OF PLANTS. 525
Eranthis hyemalis, 23, 70
ERIcACER, 114, 119, 137
Erica, 21, 73, 187, 296
cinerea, 504
hyemalis, 378, 504
multiflora, 372
Tetralix, 286, 310, 405, 406, 504
Eriobotrya japonica, 295
Erodium, 463
Ervum Lens, 20, 25, 66
Erucago, 136
Erucastrum Pollichii, 271
canariense, 280
Eryngium, 113, 368, 442
viviparum, 104
Erysimum, 136, 148
Barbarea, 252
cheiranthoides, 252
officinale, 252
Erythrochiton hypophyllanthus, 32,
174

Escholzia crocea, 250
Eucomis, 103
Eugenia Jambos, 369
Euonymus japonicus, 20
latifolius, 369
EUPHORBIACE®, 114, 150, 369, 395
Euphorbia, 114, 150, 365, 371, 395
Characias, 20
Cyparissias, 20, 244
Esula, 198, 307, 310
exigua, 20
geniculata, 255, 256
helioscopia, 56
Lathyris, 244.
rosea, 369
palustris, 106, 299
Peplus, 162
pusilla, 244.
segetalis, 281

F.

Faba, see Vicia
vulgaris, 397
Fabiana, 237
Fagus silvatica, 65, 66, 197, 318, 370,
459
Festuca, 115
nemcralis, 169
ovina, 169
Ficaria, (see Ranunculus Ficaria)
ranunculoides, 70, 295, 5G0
FICOIDER, 43
Ficus Carica, 114, 204, 435
stipulata, 332
Filago, 114
germanica, 108

524:

FInicks, 21, 190, 447
Fourcroya, 115
Fragaria, xxxvi adnot., 250, 295
alpina, 271
botryformis, 47
monophylla, 396
vesca, 20, 70, 116, 195, 275, 281,
406, 503
Fraxinus excelsior, 13, 21, 66, 325, 396,
421
Ornus, 21
Fritillaria imperialis, 21, 45, 296, 462,
506

INDEX OF

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

G.

Gagea, 365, 367
arvensis, 375, 508
Gaillardia, 269, 271
Galanthus, 296
nivalis, 300, 506
Galeobdolon luteum, 226, 238
Galeopsis, 429
Ladanum, 238
ochroleuca, 44
Tetrahit, 429
Galium Aparine, 205, 325
Mollugo, 321, 325
verum, 325
Gaudichaudiex, 334
Gardenia, 296, 377
florida, 504
Fortuniana, 604
radicans, 504
Gaura biennis, 20
Genista, 295
tinctoria, 502
sibirica, 502
Scoparia, 502
GENTIANACE®, 137, 150, 505
Gentiana, 71,73, 137, 150, 252, 296,
Amarella, 166, 305, 310, 371, 505
campestris, 250, 299, 44.2
purpurea, 389
GERANIACE®, 113, 1387, 501
Geranium, 20, 137, 221, 246, 250, 252,
292, 295, 418
columbinum, 461
nodosum, 34, 65
pratense, 501
sylvaticuin, 501

NAMES OF PLANTS.

GESNERACES, 38, 114, 222, 505
Gesnera, 33, 44, 95, 171, 339, 357, 427
Geroldtiana, 88, 89
zebrina, 355
Geum, 121, 137, 465
coccineum, 275
rivale, 122,130, 131, 250, 252, 281,
503
Gilia capitata, 281
glomeruliflora, 253, 271
Gladiolus, 21, 296
tristis, 506
Glaucium luteum, 66
Gleditschia, 30, 177, 364.
triacanthos, 44, 48
Glochidion, 310
Gloxinia, 171, 206, 207, 222, 226, 238,
284, 291, 296, 365, 451, 506
Giyceria aquatica, 169
fluitans, 169, 278
Godetia, 295, 510
Godoya, 374
Gomphia, 281
Gongora, 35
Goodenia ovata, 21, 31
GRAMINACE®, 115, 138, 278, 350, 391
Gratiola, 296
Guarea, 508
Gypsophila, 149

H.

Habenaria, 138, 238
chlorantha, 382
Halenia, 222
heterantha, 222,
Hedera Helix, 65
Hedypnois, 114
Helianthemum, 132, 137, 295, 404.
vulgare, 501
Helianthus, 38, 44, 66
annuus, 20
tuberosus, 20
Helleborus, 23. 288
foetidns, 4.4.2
olympicus, 284
Heliotropium peruvianum, 510
Helwingia, 174.
Hemerocallis, 138, 296, 310, 507
disticha, 507
fulva, 507
Hepatica, 295, 463
triloba, 500
Heracleum, 82, 113, 137, 150, 262, 365.
368
Sphondylium, 252, 256, 339
Hermesia castaneifolia, 194
Herreria parviflora, 141, 150

226

INDEX OF

Hesperis, 19, 136, 295
matronalis, 20, 252, 280, 325, 500
Heterocentron, 354
Hibiscus, 137, 298, 295,
albus, 501
flavescens, 501
Rosa sinensis, 501
Syriacus, 20, 501
Hieracium, 138
aureum, 17
prealtum, 339
Pilosella, 20
umbellatum, 20
HrProcasTaNne®, 501
Hipppeastrum, 296
equestre, 506
Hippuris, 196
vulgaris, 325, 406
Hodgsonia, 326, 474
Holcus mollis, 169
Honckenya peploides, 196, 406, 461
Hordeum, 115, 351
nepalense, 174, 175
trifurcatum, 174, 175
Humulus Lupulus, 1938, 244, 435, 472
Hyacinthus, 138, 150, 262, 296, 360,
361, 486 adnot
comosus, 409
orientalis, 21, 45, 48, 54, 172, 189,
286, 299, 348, 507
Pouzolzii, 170
Hydrangea, 417
Hydrocera triflora, 359
HYDROCHARIDACE®, 506
Hydrocharis, 296
morsus rane, 506
Hydrocotyle, 113
HYDROPHYLLACE®, 138
Hydrophyllum, 138
Hymenocallis, 404
americana, 462, 463
Hyoscyamus, 430
Hypericum, 442
perforatum, 369, 458
Hypnum triquetrum, 49
Hypocheris, 73, 138
radicata, 250, 339, 437, 457
Hyssopus officinalis, 20, 325

297, 360, 510

if
Tberis, 295, 364
amara, 500
umbellata, 500

Ilex Aquifolium, 21, 66, 447
Impatiens, 161, 231, 295, 299
Balsamina, 238, 502

Imperatoria, 82

NAMES OF PLANTS. 525

Indigofera, 459
Inula, 20
Ionopsidium acaule, 362
Ipomoea, 296, 510
pandurata, 505
IrIpAcEm, 138, 506
Irina, 64
glabra, 65
Iris, 138, 286, 296, 859, 360, 361, 365,
401, 430
Kempferi, 506
sibirica, 506
versicolor, 45
Tsochilus, 386
Ixia carminosa, 84
Ixora, 296
grandiflora, 504

J.

Jasione, 20, 114

JASMINACE®, 137, 504

Jasminum, 137, 296, 360, 400
grandiflorum, 288, 505
hirsutum, 505
nudiflorum, 21
officinale, 21, 505
Sambae, 505

Jatropha Pohliana, 254, 256

Juglans, 244, 400
nigra, 396
regia, 66, 195

JUNCACER, 115, 167, 169

Juncus, 115, 317
articulatus, 431
conglomeratus, 325
uliginosus, 107

Juniperus virginiana, 194
sinensis, 217

Jussieua, 81, 180

Justicia oxyphylla, 25

K.

Kerria, 295
Knautia arvensis, 20, 114
Kochia Scoparia, 430

L.

LABIATH, 138, 429
Laburnum (see Cytisus), 65, 157, 189,
222, 226
Lacistema, 359
Lactuca, 114
sativa, 11, 20, 33, 44, 313

526 INDEX OF
Lambertia, 3865
Lampsana, 114
Lamium, 73, 238, 361
album, 62, 63, 66, 86, 409
amplexicaule, 404:
purpureum, 66, 325, 4.04.
Larix, 114
europea, 21, 90, 245, 435
microcarpa, 192
Lathyrus latifolius, 262
tuberosus, 30
Laurus, 296, 362
nobilis, 506
Sassafras, 250, 331, 506
Lavatera trimestris, 20
Lebeckea, 459
Lecythis, 149, 180
LEGUMINOS#, 48, 71, 73, 106, 113, 122,
137, 139, 146, 147, 149, 272,
276, 429, 434, 444,
Leitnera floridana, 194.
Leontice, 125
Leontodon, 17, 20, 44, 163, 243, 442
Lepidium, 148, 364.
sativum, 57
Lepyrodia hermaphrodita, 197
Leskea sericea, 49
Leucanthemum, 86
Leucobryum giganteum, 194
Leucoium, 150, 296
eestivum, 84, 138, 350
vernum, 350, 506
Lilium, 73, 106, 115, 296, 367, 375,
421
auratum, 73, 289, 400
bulbiferum, 45
candidum, 21, 286, 325, 375, 507
cruentum, 21
lancifolium, 35, 400, 443
longiflorum, 310
Martagon, 21, 286, 325,
tigrinum, 306, 310
Linaria, 137, 229, 230, 233,
361, 365, 367, 405
zruginea, 238
chalepensis, 238
Cymbalaria, 238
decumbens, 238
Elatine, 238
origanifolia, 238
Pelisseriana, 238
pilosa, 238
purpurea, 20, 44, 238
spuria, 238
triphylla, 238
vulgaris, 162, 226, 234, 235, 238
316, 374, 505
triornithophora, 238
Linum, 335

507

296, 316,

NAMES OF

PLANTS.

Linum usitatissimum, 20

Liquidambar, 362

Listera ovata, 398

LOBELIACE®, 72, 114

Lobelia, 211, 424

Lolium, 86, 113, 115, 351
perenne, 61, 169, 325

Lonicera, 38, 44, 73, 82, 226, 271, 281,

296, 297
brachypoda, 358
Caprifolium, 408
Periclymenum, 66, 226, 251, 256,
262, 263, 338, 379, 404, 406, 504.

Xylosteum, 252, 358

Lopezia, 298, 410

Lotus, 113, 295, 360, 510
corniculatus, 104, 377, 436, 502
uliginosus, 363

Lowea, 396

Lunaria, 136, 364

Lupinus, 106, 165, 226, 280
polyphyllus, 238

Lycaste Skinneri, 93, 95, 291

Lychnis, 113; 137, 148, 295
chaleedonica, 501
Coronaria, 107, 510
dioica, 67, 252, 262, 280, 404, 464
flos cuculi, 501
sylvestris, 252, 501
vespertina, 501
Viscaria, 501

Lycium, 365, 367
europeum, 250

Lycopersicum, see Solanum
esculentum, 389

Lysimachia, 119
Ephemerum, 271, 281
nummularia, 505
vulgaris, 20, 87

Lythrum, 335
Saliearia, 374

M.

Melenia, 403
Mesa, 145, 371
MAGNOLIACEM, 122
Magnolia, 288, 388, 440

Campbelli, 427

fuseata, 304, 310, 427
MALPIGHIACE®, 137, 334, 403, 404, 406
Malus, 78, 79, 388, 389, see Pyrus
MALvVACEs, 137, 149, 288, 292, 295,

362, 395

Malva, 295

crispa, 448

moschata, 501

rotundifolia, 501

sylvestris, 252

INDEX OF

Mangifera, 159, 869
Marchantia, 17+
Maregraavia, 23
umbellata, 38382
Masdevallia, 27, 95
Mathiola, 136, 148, 295
annua, 361, 500

incana, 20, 38, 44, 68, 299, 500

glabrata, 500
Matricaria Parthenium, 28]
Maxillaria, 383

Medicago, 137, 149, 262, 295, 364, 50

lupulina, 404, 432
maculata, 218, 271
Melastoma, 74, 125, 424
Melia Azederach, 21
Melianthus major, 298, 407
Melilotus, 137, 139, 147, 149, 262
arvensis, 374
leucantha, 432
macrorhiza, 271
officinalis, 404
Melittis, 361
Mentha, 238, 361, 459
aquatica, 325
viridis, 325
Merenrialis, 62, 194, 365
annua, 198
perennis, 66
Mesembryanthemum, 26
Metrosideros, 103
Miconia, 355
Mimosa, 365
Lophantha, 31
Mimulus, 73, 284, 296
luteus, 505
Mnium serratum, 49
Mirabilis, 296, 418
Jalapa, 506
Modecea, 326
Mollugo Cerviana, 398
Momordica Elaterium, 20
Monarda fistulosa, 298
Morus, 193, 354, 459
Mozinna, 399
peltata, 194
Musa, 245, 407
Muscari, 408, 461
comosum, 340, 467
Muscr, 174
Musseenda, 249, 283, 429
Myosotis cxspitosa, 281
palustris, 375
scorpioides, 20
Myosurus, 440
Myristica moschata, 194
Myrtacem, 80, 149, 362, 395
Myrtus, 295, 310
communis, 503

NAMBPS OF PLANTS. Jal

Narcissus, 21, 67, 138, 286, 296, 360,

365, 443

aureus, 506

biflorus, 34, 45, 506

chrysanthus, 38, 45

concolor, 506

Cypri, 506

Jonquilla, 506

incomparabilis, 38, 45, 301, 506

italicus, 506

lobularis, 506

major, 154

montanus, 301

poculiformis, 506

poeticus, 24, 33, 301, 506

pseudo-narcissus, 506

Tazetta, 45, 300, 506

Telamonius, 506

| Nasturtium, 136, 271

amphibium, 181
Nelumbium, 295
speciosum, 500
Nepeta diffusa, 238
Nepenthes, 328, 473
Nephrodium molle, 447
Nerium Oleander, 62, 66, 296, 301, 377
odorum, 505
Neuropeltis, 32
Nicandra physaloides, 458
Nicotiana, 23, 31, 73
rustica, 281

| Nigella, 220, 262, 295, 297, 364, 374,

399
damascena, 226, 269, 271, 286, 375,
500
Nuytsia, 371
NYMPH HACER, 148
Nymphiea, 50, 141, 143, 148, 285, 295
alba, 162
dentata, 256, 261
euineensis, 170
Lotus, 277, 280

O.

Octadenia, 364
Odontoglossum Alexandre, 387, 403
Uro Skinneri, 463
(Enanthe, 82, 114, 360, 365, 457
crocata, 80, 359, 457
(Enothera, 44, 367
striata, 252
Olea europeea, 21, 157, 420
ONAGRACER, 81, 137, 406, 503
Oncidium, 67, 114, 227
abortivum, 462
bicolor, 45
Cebolleta, 107

528

Oncidium cucullatum, 91
heteranthum, 224
ornithorhyncum, 45

Ononis minutissima, 404:
monophylla, 396

Opercularia, 38

Ophrys, 114:
apifera, 360, 398
aranifera, 35, 42, 45, 238, 298,

383, 384, 385, 386, 398
insectifera, 27, 301, 382
fucifera, 506

Opuntia, 81, 113, 149, 180
fragilis, 107, 178
monacantha, 178
Salmiana, 107, 178, 179

ORCHIDACER, 27, 34, 42, 91, 112, 114,

128, 188, 150, 1538, 209, 228,
290, 345, 360, 366, 367, 380—
387, 397

Orchis, 138, 150, 153, 154, 227, 296
conopsea, 238
latifolia, 238
mascula, 153, 154, 238, 299, 387,

506
militaris, 387
Morio, 238, 382, 384, 387, 506
palustris, 366
papilionacea, 238
pyramidalis, 128, 238, 506
sambucina, 63, 66
simia, 238

Ornithogalum, 114, 360, 365, 367
longebracteatum, 171

OROBANCHACE®, 137

Orobanche, 73, 137, 209, 296, 505
gracilis, 442

Orobus, 295
vernus, 502
viscosus, 502

Ouvirandra, 458

OXALIDACER, 403

Oxalis, 295, 329, 404, 459
Acetosella, 404
carnea, 502

ips

Peonia, 295, 300, 364, 399, 407
Moutan, 500
officinalis, 261
paradoxa, 500
tenuiflora, 500

PALMACER, 155

Panax, 331

Papaver, 289, 295, 297, 473
bracteatum, 28, 304, 310, 500
nudicaule, 310
orientale, 250

INDEX OF NAMES

OF PLANTS.
Papaver Rheeas, 429, 458, 500
somniferum, 28, 305, 310, 500
PAPAVERACER, 99, 500
PAPILIONACER, 897, 502, see Legumi-
NOSE
Paris quadrifolia, 31, 358, 360, 367, 396
Paritium, 137
Parnassia, 364
Paronychia serpyllifolia, 473
Parthenium inodorum, 243
Passiflora, 137, 201, 301, 365, 463, 473
coerulea, 185
palmata, 185
quadrangularis, 181
PASSIFLORACE®, 137
Pastinaca, 82
sativa, 272
Pavia, 137
Pedicularis, 238
euphrasioides, 238
sylvatica, 44, 223, 226, 238
Peganum, 363
Pelargonium, 22, 23, 25, 63, 107, 113,
208, 221, 225, 295, 336, 360,
373, 389, 418, 419, 434:
grandiflorum, 86
inquinans, 30, 226
zonale, 106, 226, 501
Peltaria, 137
alliacea, 250, 251, 252
Pentstemon, 226, 455
Pereskia, 107, 118, 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

Phalenopsis, 238
amabilis, 227
equestris, 231, 238
Schilleriana, 224
Pharbitis, 73
Phaseolus, 21, 298, 364
PHILADELPHACE®, 113, 187, 149
Philadelphus, 118, 1387, 142, 149, 295,
364, 367
coronarius, 503
speciosus, 178, 298
Philyra brasiliensis, 198
Phleum, 115, 188
phalaroides, 169
pratense, 325
Phlomis, 138
biloba, 72
fruticosa, 119, 238

INDEX OF

Phlox, 73
Pheenix dactylifera, 318
Phylica, 325
Phyllanthus longifolius, 198
Phyllarthron, 64, 396
Phyteuma odoratum, 271
orbiculare, 20
spicatum, 166, 252
Phytolacca, 20
Picea, see Abies
Lowii, 203
nobilis, 203
Nordmanniana, 203
Webbiana, 203
Picrasma, 509
Pimpinella magna, 66
Saxifragra, 66
Pinckneya, 429
Pinus, 11, 19, 114, 318, 346, 471, (see
Abies, Larix, Cedrus)
alba, 192
pinea, 90, 218
Pinaster, 13, 21
sylvestris, 21, 349
Pistacia Lentiscus, 197
Pisum, 137, 295
sativum, 27, 30, 31, 95, 166, 302,
372, 432, 472, 502, 510
Platycodon, 296
grandiflorum, 504:
PLANTAGINACE®, 114
Plantago, 114, 136
Coronopus, 458
lanceolata, 61, 104, 108, 110, 111,
243
media, 20, 111
major, 65, 108, 109, 111, 243, 372,
374
maritima, 108
Platanus, 50
Plectranthus fruticosus, 238
PLUMBAGINACER, 114
Plumbago, 360
Poa alpina, 169
annua, 169 +
bulbosa, 169
pratensis, 169
trivialis, 169 .
Podalyria myrtillifolia, 44
Podocarpus, 426
Podophyllum, 295
peltatum, 500
Podospermum laciniatum, 247, 250
Poggendorffia, 294
Pogonia ophioglossoides, 238, 386
Poinsettia, 340
Polemonium ceruleum, 20, 66, 253,
305, 310, 404
Polianthes, 296

NAMES

|
|

OF PLANTS.
Polygala vulgaris, 399
PoLYGONACER, 114, 138, 150
Polygonatum anceps, 42, 45
multifolium, 28, 31
Polygonum, 114, 365
orientale, 31, 390
viviparum, 106, 169
Polypodium anomalum, 190
PomacE®, 70, 71, 77, 79, 142, 405,
503
Pomaderris elliptica, 355
Pomax, 38
Populus, 202, 309
alba, 66
Portulaca, 295
Potamogeton, 329
bifolinm, 434
Potentilla, 139, 140, 149, 295, 374, 438
alpestris, 503
argentea, 280
anserina, 503
major, 374
nepalensis, 250, 252, 275, 280
reptans, 503
Tormentilla, 503
Poterium, 113
Sanguisorba, 107
polygamum, 281
Primula, 71, 73, 114, 121, 138, 150,
296, 335, 360, 365, 367
acaulis (see vulgaris), 45, 248, 250,
308, 310, 377, 504
Auricula, 45, 133, 271, 282, 504.
denticulata, 20, 504
elatior, 45, 250, 504:
calycanthema, 283
imperialis, 103
officinalis, 250, 283
prenitens, 271, 504 (see sinensis)
sinensis, 24, 45, 95, 98, 103, 133,
253, 256, 262, 263, 271, 281, 297,
814, 315, 389, 429, 441, 449
variabilis, 104
veris, 17, 20, 35, 39, 45, 105, 164
villosa, 504
vulgaris, 17, 70, 106 (see acaulis
and officinalis)
PRIMULACER, 98, 114, 118, 121, 133,
138, 150, 262, 339, 366, 504
Prismatocarpus, 114, 150, 178
Prunella vulgaris, 226
Prunus, 137, 203, 295, 300, 360, 364,
367 (see Cerasus, Amygdalus,
Persica)
Armeniaca, 44
Cerasus, 74
domestica, 366, 464
Lauro Cerasus, 21, 446
spinosa, 44
34

530 INDEX OF NAMES OF PLANTS.

Prunus sylvestris, 21
Pseudostachyum polymorphum, 168
Psittacanthus, 371
Ptelea, 364
Pterandra, 290
Pteridophyllum, 331
Pterisanthes, 328
Pteris quadriaurita, 333
Punica Granatum, 21, 295, 325
Pyrethrum, 114.
inodorum, 431
Parthenium, 339

Pyrus, 82, 107, 113, 121, 137, 149, 203
communis (Pear), 36, 66, 162, 178,

422, 423, 503
dioica, 304 U7
japonica, 161

Malus (Apple), 44, 78, 79, 166, 240,
250, 282, 295, 310, 325, 327, 375,

406, 420, 503
spectabilis, 508
torminalis, 325

Q.
(Juercus, 51

Cerris, 66
pubescens, 66

Kt.

Raphanus sativus, 161, 252, 327, 360,

369

RANUNCULACE®, 113, 122, 136, 148,

195, 246, 339, 410, 499

Restio, 115, 194
Rhamnus catharticus, 463

Frangula, 280

| Rhinanthus crista galli, 238
Rhodiola rosea, 197

Rhodora, 73

Rhododendron, 72, 73, 91, 133, 137,

176, 289, 290, 296, 453
indicum, 504
linearilobum, 72

Rhus Cotinus, 62, 66, 348, 409, 467,

472
Toxicodendron, 66

Ribes, 296

Ranunculus, 19, 55, 113, 119, 258, 288, |

295, 296, 297, 360, 367, 407
aconitifolius, 500
aeris, 67, 250, 436, 500
aquatilis, 458
asiaticus, 500

auricomus, 195, 285, 397, 406, 461,

469

bulbosus, 17, 20, 44, 195, 406, 500

bullatus, 500

Ficaria, 70, 96, 195, 261, 368, 406

fluitans, 433, 437
gramineus, 500
Lingua, 44, 67
Philonotis, 20, 339
tripartitus, 20, 46
repens, 23, 252, 261, 500
RESEDACE®, 137, 148
Reseda, 67, 137, 141, 148
lutea, 252, 271
Luteola, 61, 112, 399
odorata, 20, 44, 280
Phyteuma, 252, 261
RESTIACER, 115, 167, 189, 350, 463

nigrum, 66, 296
sanguineum, 503

Richardia zthiopica, 337, 358
Ricinus communis, 198, 300
Ricotiana, 364.

Robinia, 365

Pseudaeacia, 44, 204, 317, 325, 396

Rosacez, 70, 71,77, 80, 113, 121, 122,

137, 149, 246, 274, 405, 406, 502

Rosa, 21, 77, 79, 82, 113, 121, 130,

137, 139, 149, 151, 152, 162, 176,
184, 185, 186, 204, 246, 247, 250,
257, 258, 262, 271, 274, 288, 289,
295, 367, 368, 394, 473, 502

alpina, 209

arvensis, 307, 310, 404

Banksiwe, 502

berberifolia, 396

canina, 502

centifolia, 31, 502

Carolina, 502

cinnamomea, 502

damascena, 502

diversifolia, 280

Eglanteria, 502

gallica, 31, 502

indica, 502

moschata, 502

nivea, 502

pimpinellifolia, 502

rubiginosa, 502

spinosissima, 502

sulphurea, 502

, RUBIACER, 45, 80

Rubia tinctorum, 322, 325
Rubus, 137, 252, 279, 399, 406, 429,

465

arcticus, 67

cesius, 281, 285, 502

corylifolius, 502

fruticosus, 66, 281, 374, 380, 461
502

Idzeus, 396

rosifolius, 502

Rudbeckia, 82, 114, 127

INDEX OF

Ruellia clandestina, 403, 404
Rumex, 45, 138, 150, 325
arifolius, 278, 281
crispus, 304, 310, 448
scutatus, 278, 281, 431
Ruscus, 328, 470
aculeatus, 60, 318, 348, 445
Russellia juncea, 20
RuTAcex®X, 137, 149
Ruta, 363, 367, 371

TM

Sagina, 397
procumbens, 501
Sagittaria, 296, 329
latifolia, 507
sagittifolia, 325, 507
Salisburia adiantifolia, 61
Salix, 166, 168, 244, 310, 326, 349
babylonica, 202, 257, 262, 299, 309
326
calyculata, 29
caprea, 271, 299
cinerea, 41, 45, 189, 299
fragilis, 357
monandra, 29
nigricans, 299
pendula, 357
repens, 194
silesiaca, 299
vitellina, 21
Salpiglossis straminea, 44
Salvia, 226
officinalis, 66
pratensis, 36
Verbenaca, 87
Sambucus, 296, 359, 360, 365, 367
nigra, 21, 55, 66, 164, 325, 396
racemosa, 66
Sanguinaria, 295
canadensis, 500
Sanguisorba, 113
officinalis, 107
SANTALACE®, 122, 138, 150
SAPINDACEX, 137, 448
Saponaria, 28, 67, 76, 149, 295, 297,
300, 360, 379
officinalis, 97, 301, 501
Sarothamnus, 360, 363
Scoparinus, 363
Saxifraga, 33, 44, 106, 113, 296, 390
crassifolia, 307, 310
decipiens, 288
foliosa, 281
granulata, 298, 504
irrigua, 20
longifolia, 404

NAMES OF

PLANTS. 531
Saxifraga mutata, 20
SAXIFRAGACE®, 80, 81
Scabiosa, 35, 106, 114, 138, 141, 296,
297, 429
agrestis, 252
arvensis, 325
atropurpurea, 62, 66
Columbaria, 252, 281
Seandix, 125
Schismatopera distichophylla, 194
Schcenodorus, 364 adnot
Scheenus cephalotes, 245
Scheenodon, 365
Sciadopitys verticillata, 352
Scilla, 296
autumnalis, 507
nutans, 507
Scirpus lacustris, 325
Scolopendrium D‘Urvillei, 332
vulgare, 64, 314, 326, 328, 459, 467
Scorzonera, 19
octangularis, 247, 250
Scrophularia, 114, 455, 463
aquatica, 226, 238, 262, 28
arguta, 334
nodosa, 44, 106, 281
ScROPHULARIACER, 114, 121, 137, 150,
222, 505
Secale cereale, 55, 115
Sechium edule, 81, 202
Sedum cristatum, 18, 20
reflexum, 20
Selenipedium caudatum, 224, 227

| Selinum, 82, 114

caruifolium, 281
Sempervivum, 105, 113, 204, 509
montanum, 310
sediforme, 164.
tectorum, 292 adnot, 308, 310
Senecio, 44, 114
vulgaris, 247, 250, 252, 339, 432,
437, 439, 457
Serissa, 296, 377, 504
Sesamum, 365
indicum, 238
Seseli, 113, 281
coloratum, 166
| Sideritis, 238, 365

| Silaus, 113, 365

|
|

|
|

Silene, 44, 137, 148, 295, 301
conica, 68
Otites, 464
inflata, 501

| Sinapis, 57, 136, 270, 295, 371

arvensis, 181, 250, 261, 264, 271,
280, 501
Sinningia purpurea, 431
Sisymbrium, 136
officinale, 250, 261, 271, 280

532

Sisymbrium tenuifolium, 252
Slateria, '74
SOLANACER, 150, 4.30, 505
Solanum, 73, 125, 150, 296, 360, 365,
367, 370, 424, 453
amazonicum, 430
Dulcamara, 66, 288, 430, 504
esculentum, 74
cornutum, 430
Lycopersicum, 38, 44, 74, 171, 391,
44,2
tridynamum, 430
tuberosum, 54, 142, 288, 333, 420
Vespertilio, 430
Sophora, 55
Spartianthus, 295
junceus, 502
Spartium, 295
junceum, 18, 21
Scoparium, 21
Spathiphyllum, 245
Spilanthes, 138, 365
oleracea, 44
Spinacia oleracea, 31, 197
Spirea, 21, 137, 295
Filipendula, 503
obiongifolia, 252
prunifolia, 503
Reevesii, 503
strobilacea, 503
Ulmaria, 503
Spiranthes, 319
Splachnum vasculosum, 49
Stachys, 138, 339, adnot
germanica, 310
lanata, 375
sylvatica, 74, 253, 271, 281
Stackhousia juncea, 334
Stangeria paradoxa, 172
Stapelia, 20
Staphylea pinnata, 30
Stauntonia latifolia, 194
Stellaria, 148, 404, 406
media, 271, 279, 280
STELLAT®, 396, 408
STERCULIACE®, 362, 464
Sterculia platanifolia, 20, 256
Sternbergia, 296
lutea, 506
Stratiotes aloides, 406
Strelitzia juncea, 459
regina, 25
Streptocarpus Rexii, 15, 44, 226, 227,
461
Strophanthus, 326
Suaeda, 365
fruticosa, 430
maritima, 20, 21
Suregada, 198

INDEX OF NAMES OF PLANTS.

| Symphoricarpus racemosus, 66

Symphyomyrtus, 38

Symphytum, 71, 138, 365
officinale, 253, 262, 263, 271
Zeyheri, 271

| Syringa, 296, 360, 367

persica, 44, 61, 66, 284, 505

vulgaris, 76, 505

AN

Tabernemontana, 296
coronaria, 504
Tacsonia pinnatistipula, 294
Tamus communis, 21
Taraxacum, 164 (see Leontodon)
Taxodium, 114
distichum, 444
Taxus baccata, 11, 21, 90
Tetragonia, 113
expansa, 142, 180
TETRAGONIACE®, 149
Teucrium campanulatum, 226, 233, 238
Thalictrum, 295, 500
minus, 307, 310
Thea, 295, 502
Thelymitra, 224.
Thesium, 121, 123, 138, 150
Thlaspi arvense, 250, 271
bursa-pastoris, 252, 428
Thuja occidentalis, 319
orientalis, 21
Thymus Serpyllum, 325, 405, 406
Thysselinum, 137, 365
palustre, 437, 457
Tigridia, 360, 361, 365
Pavonia, 35, 462
Tilia asplenifolia, 66, 459
europea, 22, 30, 65
parvifolia, 66
Tiliacew, 137, 362
Tithonia, 26
Tofieldia, 296
calyculata, 300
Torenia scabra, 406
Torilis, 82, 149
Anthriscus, 256, 271, 281, 339,
406, 441, 442
Tradescantia, 245, 296, 360, 365, 367,
404

virginica, 88, 507
Tragopogon, 19, 20, 73, 114, 431
orientale, 250
pratense, 247, 250, 442, 457
Trapa natans, 202, 364
Trichostomum rigidulum, 49
Trifolium, 106, 113, 121, 187, 139, 149,
295, 360, 364

INDEX OF
Trifolium hybridum, 262, 399, 406,
4OT

pratense, 20
repens, 20, 23, 68, 70, 98, 145, 146,
226, 231, 238, 250, 252. 256,
260, 262, 265, 271, 276, 279,
356, 368, 397, 399, 406, 407,
434, 436, 438, 502
resupinatum, 20
Trillium grandiflorum, 507, 510
Trinia vulgaris, 405
Triphasia aurantiaca, 369
Triticum, 86, 115, 350, 391
repens, 106, 325
vulgare, 55, 113
Triumfetta, 137, 252, 259, 260, 262,
265, 271, 280
Trollius europeus, 66, 295, 500
TROPEOLACE®, 149, 501
Tropzolum, 149, 398, 283
majus, 20, 222, 225, 226, 232, 238,
251, 252, 254, 256, 271, 280, 295,
310, 406, 442, 501
minus, 501
Tulipa, 35, 45, 67, 75, 84, 85, 138, 262,
300, 302, 348, 359, 360, 361, 365,
367, 388, 390, 421
Gesneriana, 31, 250, 310, 315, 442,
507
sylvestris, 507
Turritis, 271
glabra, 252
Typha, 189

Ulex, 295, 360
europeus, 377, 502
Ulmus americana, 66
campestris, 31, 52, 62, 157, 325,
353, 427
UMBELLIFER®, 37, 71,73, 80,107, 113,

NAMES OF

PLANTS. 533
Valeriana, 114, 165, 419
dioica, 325
officinalis, 325
Valisneria spiralis, 319, 433
Verbascum, 73, 116, 137, 225, 253, 430
australe, 35
nigrum, 226, 251
phlomoides, 281
Thapsus, 404
Verbena, 67, 68, 506
Veronica, 60, 150, 296, 334, 360, 361,
365, 375, 505
austriaca, 66
Beccabunga, 468
Chameedrys, 442
latifolia, 459
longifolia, 325
spicata, 325, 374

| Viburnum, 44, 296, 417

Opulus, 473
Vicia, 30, 365, 369, 472
Faba, 434.
Vinca, 137, 296
herbacea, 389
major, 505
minor, 44, 99, 358, 390, 505
rosea, 31
Viola, 23, 137, 225, 229, 289, 295, 297,
405
grandiflora, 501
hirta, 226, 238
odorata, 20, 94, 220, 226, 238, 286,
404, 429, 501, 508
tricolor, 461, 501
VIOLACER, 137, 334, 403, 406, 500
Viscum album, 56, 369, 509

| VITAcE®, 137
| Vitex incisa, 238
| Vitis, 417

121, 127, 131, 132, 137, 139, 140, |

143, 149, 150, 162, 244, 257, 261,
339, 368, 395, 405, 406, 407, 437,
439

Uredo candida, 279

URTICACER®, 114

Urtica dioica, 62, 66, 194.

Usteria, 283

We

Vaccinium, 68
Valantia cruciata, 44.
VALERIANACER, 114

vinifera, 20, 29, 34, 44, 66,137, 157,
182, 183, 211, 280, 374, 422 424

W.

Wedelia perfoliata, 442
Weigela rosea, 358

Welwitschia, 162, adnot

Wildenovia, 115, 167, 168
Wistaria, 364
sinensis, 226

X.

Xanthosoma appendiculatum, 31
Xanthoxylum, 21 j
Xylophylla, 328

By)

534 INDEX TO NAMES OF PLANTS.
YA Zea Mays, 21, 113, 136, 191, 197, 300,
Y 361 310, 350, 369, 466
brag | Zingiber Zerumbet, 224, 227
flexilis, 361 Zinni
innia, 44
7 elegans, 20

| Zygopetalum maxillare, 398
Zamia, 170

ERRATA.

The reader is requested to make the following corrections :—
Page 182. Fig. 94 should be 94*.

Page 194. The reference 3 applies not to the nutmeg but to the hop,
figured at p. 193.

Page 309. Fig. 165 legend—for Sempervivun tecotorum read Semper-
vivum tectorum.

PRINTED BY J, E. ADLARD, BARTHOLOMEW CLOSE,

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