LANiJSCAKt
ARCHITECTURE
Ex
Libris
BEATRIX
JONES
REEF POINT GARDENS
LIBRARY
The Gift of Beatrix Farrand
to the General Library
University of Calif ornia,Berkeley
GRAY'S BOTANICAL TEXT-BOOK.
f*
VOLUME I.
STRUCTURAL BOTANY.
GRAY'S BOTANICAL TEXT-BOOK
CONSISTS OF
VOL. I. STRUCTURAL BOTANY. By ASA GRAY.
II. PHYSIOLOGICAL BOTANY. By GEORGE L. GOODALE.
III. INTRODUCTION TO CRYPTOGAMIC BOTANY, BOTH
STRUCTURAL AND SYSTEMATIC. By WILLIAM G.
FARLOW. {In preparation.)
IV. SKETCH OF THE NATURAL ORDERS OF PH^ENOGAMOUS
PLANTS; their Special Morphology, Useful Pro-
ducts, &c. {In preparation.)
GRAY'S BOTANICAL TEXT-BOOK.
(SIXTH EDITION.)
VOL. I.
STRUCTURAL BOTANY
OR
ORGANOGRAPHY ON THE BASIS OF
MORPHOLOGY.
TO WHICH IS ADDED THE PRINCIPLES OP
TAXONOMY AND PHYTOGRAPHY,
*]>L-
& logsatg of Botanical
BY
ASA, GRAY, LL.D., ETC.,
FISHER PROFESSOR OF NATURAL HISTORY (BOTANY) IN
HARVARD UNIVERSITY-
NEW YORK : CINCINNATI . : . CHICAGO
AMERICAN BOOK COMPANY
FROM THE PRESS OF
IVISON, Bl.AEKMAN & COMPANY
Copyright,
BY ASA GRAY.
187.
idd'l
Farrand Gift
LANDSCAPE
ARCK.
LIBRARY
PREFACE.
THE first edition of this treatise was published in the
year 1842, the fifth in 1857. Each edition has been in
good part rewritten, the present one entirely so, and the
compass of the work is now extended. More elementary
works than this, such as the author's First Lessons in
Botany (which contains all that is necessary to the prac-
tical study of systematic Phtenogamous Botany by means
of Manuals and local Floras), are best adapted to the
needs of the young beginner, and of those who do not
intend to study Botany comprehensively and thoroughly.
The present treatise is intended to serve as a text-book for
the higher and completer instruction. To secure the
requisite fulness of treatment of the whole range of sub-
jects, it has been decided to divide the work into distinct
volumes, each a treatise by itself, which may be indepen-
dently used, while the whole will compose a comprehensive
botanical course. This volume, on the Structural and
Morphological Botany of Phgenogamous Plants, properly
comes first. It should thoroughly equip a botanist for the
scientific prosecution of Systematic Botany, and furnish
needful preparation to those who proceed to the study of
Vegetable Physiology and Anatomy, and to the wide and
varied department of Cryptogamic Botany.
212
IV PREFACE.
The preparation of the volume upon Physiological
Botany (Vegetable Histology and Physiology) is assigned
to the author's colleague, Professor GOODALE.
The Introduction to Cryptogamous Botany, both structu-
ral and systematic, is assigned to his colleague, Professor
FABLOW.
A fourth volume, a sketch of the Natural Orders of
Phaenogamous Plants, and of their special Morphology,
Classification, Distribution, Products, &c., will be needed
to complete the series : this the present author may
rather hope than expect himself to draw up.
ASA GRAY.
HERBARIUM OF HARVARD UNIVERSITY,
CAMBRIDGE April 10, 1879.
into sentences or appended to them, are references to the numbered para-
graphs in which the topic is treated or the term explained.
CONTENTS.
PAGE
INTRODUCTION. THE DEPARTMENTS OF THE SCIENCE .... 1
CHAPTER I. OUTLINES OF THE GENERAL MORPHOLOGY
OF PHJENOGAMOUS PLANTS 6
CHAPTER II. MORPHOLOGY AND DEVELOPMENT OF THE
EMBRYO AND SEEDLING 9
The Embryo, its Nature, Structure, and Parts 9
Development of the Dicotyledonous Embryo in Maple ... 10
In Ipomcea, or Morning Glory, &c., with Albuminous Seeds . 13
In Embryos with thickened Cotyledons 16
As of Almond, Beech, Bean, &c 17
With Hypogaeous Germination and no Elongation of Caulicle 19
In Megarrhiza, &c., with concreted Petioles to the Cotyledons 21
In Ipomoea leptophylla with foliaceous and long-petioled
Cotyledons and no elongation of Caulicle 22
In Pumpkin, &c., with no Primary Root 22
The Polycotyledonous Embryo 23
The Monocotyledonous Embryo of Iris, Onion, Cereal Grains 24
Pseudo-monocotyledonous and Acotyledonous Embryo ... 26
Dicotyledonous and Monocotyledonous Plants 27
CHAPTER III. MORPHOLOGY AND STRUCTURE OF THE
ORGANS OF THE PLANT IN VEGETATION . . 27
SECTION I. OF THE ROOT 27
Nature, Growth, and Composition 28
Root-hairs 29
Kinds of Roots 29
Duration ; Annuals 30
Biennials 31
Perennials 32
Aerial Roots 33
Epiphytes or Air-plants 35
Parasitic Plants, Green and Colored 36
VI CONTENTS.
SECTION II. OF Bui>s 40
Scaly Buds and Bud-scales 40
Naked, Subpetiolar, and Fleshy Buds 41
Bud-propagation 43
Normal, Accessory, and Adventitious Buds 44
SECTION III. OF THE STEM . 45
1. GENERAL CHARACTERISTICS AND GROWTH 45
Development and Structure 46
Ramification, Branches 47
Excurrent and Deliquescent Stems 48
Definite and Indefinite Annual Growth . 49
2. FORMS OF STEM AND BRANCHES 50
Herbs, Shrubs, Trees, Culm, Caudex, Scape 50
Climbing Stems, Twining or otherwise 51
Leaf-climbers, Tendril-climbers, and Root-climbers .... 52
Suckers, Stolons, Offsets, Runners 53
Tendrils formed of Stems 54
Sympodial and Monopodial Stems 55
Spines or Thorns and Subterranean Stems 56
Rhizoma or Rootstock 57
Tuber, Tubercles 59
Corm or Solid Bulb 61
Bulb, Bulblets 62
Condensed Aerial Stems 64
Stems serving for Foliage, Phyllocladia, Cladophylla ... 65
Frondose Stems 66
3. INTERNAL STRUCTURE . 67
Anatomical Elements 68
Endogenous Structure 70
Exogenous Structure ; its Beginning 73
First Year's Growth 74
Pith, Layer of Wood, &c 75
Bark, its Parts and Structure 76
Annual Increase in Diameter 78
Demarcation of Annual Layers . 79
Sap-wood and Heart-wood 80
Growth and Duration of Bark 81
Living Parts of a Tree or Shrub, Longevity 83
The Plant composite 84
SECTION IV. OF LEAVES 85
1. THEIR NATURE AND OFFICE 85
Parts of a Leaf 85
Duration, Defoliation, Normal Position ... 86
CONTENTS. Vii
2. THEIR STRUCTURE AND FORMS AS FOLIAGE 87
Internal Structure or Anatomy 87
Parenchyma-cells 88
Epidermis, Stomata or Breathing-pores 89
Framework, Venation 90
Parallel-veined or Nerved Leaves , . . 91
Reticulated or Netted-veined Leaves 92
Pinnately or Feather-veined and Palmately or Radiately
Veined 93
Forms as to Outline 94
Forms as to Extremity 96
Forms as to Margin or Special Outline and Dentation ... 97
Lobation or Segmentation 98
Number and Arrangement of Parts 99
Compound Leaves, Pinnate and Palmate or Digitate, &c. . . 100
Petiole or Leafstalk 104
Stipules, Ligule, Stipels 105
Leaves in unusual Modifications 106
Such as Inasquilateral, Connate, Perfoliate 107
Vertical and Equitant 108
Without distinction of Parts 109
Stipules serving for Blade 109
Phyllodia, or Petioles serving for Blade 110
3. LEAVES SERVING SPECIAL OFFICES 110
Utilizing Animal Matter 110
Ascidia or Pitchers Ill
Sensitive Fly-traps 113
Leaves for Storage 115
Bulb-scales and Bud-scales 116
CHAPTER IV. PHYLLOTAXY, OR LEAF-ARRANGEMENT . 119
SECTION I. DISTRIBUTION OF LEAVES ON THE STEM .... 119
Phyllotaxy either Verticillate or Alternate, Cyclical or Spiral 119
Verticillate or Cyclical Arrangement 120
Alternate or Spiral Arrangement 121
Its Modes and Laws 122
Relation of Whorls to Spirals 129
Hypothesis of the Origin of Both 130
Fascicled Leaves 131
SECTION II. DISPOSITION OF LEAVES IN THE BUD 132
Vernation and Estivation; the Modes ........ 132
Direction, Dextrorse and Sinistrorse 140
Vlll CONTENTS.
CHAPTER V. ANTHOTAXY OR INFLORESCENCE .... 141
Bracts and Bractlets and their Modifications 141
Peduncles, Pedicels, Rhachis, Receptacle 143
Position of Flower-buds, Kinds of Inflorescence 144
Indeterminate, Indefinite, or Botryose 140
Raceme, Corymb, Umbel 146
Head or Capitulum 147
Syconium or Hypanthodium 148
Spike, Spadix, Ament or Catkin 149
Panicle and other Compound Forms 150
Determinate or Cymose 151
Cyme, Glomerule, &c 152
Botryoidal Forms of Cymose Type 153
Sympodial Forms 154
Scorpioid and Helicoid, the Pleiochasium, Dichasium, and
Monochasium 155
Bostryx, Cincinnus, Rhipidium, Drepanium, &c 156
Mixed Inflorescence 158
Thyrsus, Verticillaster, &c 159
Relations of Bract, Bractlet, and Flower 160
Anterior and Posterior, or Inferior and Superior 160
Median and Transverse 160
Position of Bractlets 161
Tabular View of Inflorescence 162
CHAPTER VI. THE FLOWER 163
SECTION I. ITS NATURE, PARTS, AND METAMORPHY .... 163
Floral Envelopes, Perianth, or Perigone 164
The Parts, Calyx and Corolla 165
Androecium, Stamens 165
Gynoecium, Pistils 166
Torus or Receptacle of the Flower 167
Metamorphosis 167
Unity of Type illustrated by Position and Transitions . . . 169
Teratological Transitions and Changes 170
SECTION II. FLORAL SYMMETRY 174
Symmetrical, Regular, and Complete Flower 175
Numerical Ground-plan 176
Pattern Flowers 176
Diplostemonous Type 177
SECTION III. VARIOUS MODIFICATIONS or THE FLOWER . . . 179
'l. ENUMERATION OF THE KINDS 179
2. REGULAR UNION OF SIMILAR PARTS 180
Coalescence or Cohesion 180
CONTENTS. ix
3. UNION OF DISSIMILAR OR SUCCESSIVE PARTS .... 181
Adnation or Connation 182
Hypogynous, Perigynous, Epigynous 183
4 IRREGULARITY OF SIMILAR PARTS 184
5. DISAPPEARANCE OR OBLITERATION OF PARTS .... 187
Abortion or Suppression of Parts of a Circle 187
Abortion or Suppression of whole Circles 190'
Terms therewith connected ." 191
Suppressed Perianth 191
Suppressed Androecium or Gynoecium 193
Along with suppressed Perianth 194
Neutral Flowers " . . 195
6. INTERRUPTION OF NORMAL ALTERNATION 195
Anteposition or Superposition 195
In Appearance only 196
Superposition by Spirals 196
Anteposition with Isostemony and Diplostemony 197
With Obdiplostemony 198
7. INCREASED NUMBER OF PARTS 200
Regular Multiplication 200
Parapetalous Multiplication 201
Chorisis or Deduplication 202
8. OUTGROWTHS 209
Their relation to Chorisis : Trichomes 209
Corona or Crown 210
Ligule 211
9. FORMS OF THE TORUS OR RECEPTACLE 211
Stipe, Thecaphore, Gynophore, Carpophore, &c 212
Disk 213
Hypanthium 214
SECTION IV. ADAPTATIONS OF THE FLOWER TO THE ACT OF
FERTILIZATION 216
1. IN GENERAL 215
Close and Cross Fertilization, or Autogamy and Allogamy . 216
2. ADAPTATIONS FOR ALLOGAMY OR INTERCROSSING . . . 216
Wind-fertilizable or Anemophilous Flowers ....... 217
Insect-fertilizable or Entomophilous Flowers 218
Irregularity as related to Allogamy 219
CONTENTS.
Dichogamy, either Proterandrous or Proterogynous .... 219
Proterogyny . 219
Proterandry 220
Particular Adaptations in Papilionaceous Flowers .... 225
In Kalmia-blossoms, Iris, &c 229
Transportation of Pollinia 230
In Orchidaceae and Asclepiadaeese 231
Heterogonous Dimorphism and Trimorphism 234
3. ADAPTATIONS FOR CLOSE FERTILIZATION 240
Cleistogamy 241
SECTION V. THE PERIANTH, OR THE CALYX AND COROLLA IN
PARTICULAR 243
Perianth as to Duration, Numerical Terms, Union, &c. . . . 243
Parts of Petals and of Gamophyllous Perianth 245
Forms of Corolla and Calyx 246
SECTION VI. THE ANDRGECIUM, OR STAMENS IN PARTICULAR . 249
The Stamen as a whole ; Numerical Terms 249
The Filament and the Anther ; their Modifications .... 251
Pollen 256
Pollen-tubes 258
SECTION VII. THE PISTILS, OR GYNOSCIUM 259
1. IN ANGIOSPERMS 259
Carpel or Carpophyll 260
Ventral and Dorsal Sutures ; Placenta 261
Simple or Apocarpous Pistils 262
Compound or Syncarpous Pistil 263
With two or more Cells and Axile Placentae ; Partitions . . 264
With one Cell and Parietal Placentae 265
With one Cell and Free Central Placenta 266
Anomalous Placentation 267
2. IN GYMNOSPERMS 268
Structure in Gnetaceae 269
Structure in Coniferae 270
In the Yew Family 271
In the Pine Tribe, &c 272
In the Cypress Tribe 273
Structure in Cycadaceas 274
SECTION VIII. THE OVULE 276
Its Structure and Position 277
Its Forms, Orthotropous, Campylotropous, Amphitropous,
Anatropous 278
Origin and Morphological Nature of the Ovule 282
Origination of the Embryo 283
CONTENTS. "xi
CHAPTER VII. THE FRUIT 285
SECTION I. ITS STRUCTURE, TRANSFORMATIONS, AND DEHIS-
CENCE 285
Pericarp, its Alterations, Accessions, and Transformations . 287
Dehiscence 288
SECTION II. THE KINDS OF FRUIT 291
Simple Fruits 291
Dehiscent Fruits, Follicle, Legume, Capsule, Pyxis, Silique . 292
Indehiscent Dry Fruits, Samara, Akene, Utricle, Caryopsis,
Nut, &c 294
Fleshy Fruits, Drupe, Pome, Pepo, Berry, &c 297
Aggregate Fruits 299
Accessory or Anthocarpous Fruits 300
Multiple or Collective Fruits, Syconium, Strobile, &c. . . . 301
Table of Simple Fruits 304
CHAPTER VIII. THE SEED 305
Its Stalk, Coats, and Appendages 306
Aril or Arillus 308
Nucleus or Kernel, Albumen 309
The Embryo, its Parts and Positions 311
The Cotyledons as to Adjustment arid Number 313
CHAPTER IX. TAXONOMY 315
SECTION I. THE PRINCIPLES OF CLASSIFICATION IN NATURAL
HISTORY 315
Individuals 315
Species 317
Varieties, Races, &c .' 318
Cross-breeds and Hybrids 321
Genera 323
Orders, Classes, Tribes, &c 325
Sequence of the Grades 327
Nature and Meaning of Affinity 327
Theory of Descent and Natural Selection 328
SECTION II. BOTANICAL CLASSIFICATION 331
Ante-Linnaean Classifications 332
Linnaean Classification 333
Sexual Artificial System 334
Natural System 338
As presented by Jussieu 339
Some of its Modifications , . 340
Xil CONTENTS.
CHAPTER X. PHYTOGRAPHY 345
SECTION I. NOMENCLATURE 345
Names of Plants, Binomial Nomenclature 346
Rules for naming Plants 347
Names of Genera 348
Names of Species, Varieties, &c 350
The Fixation, Precision, and Citation of Names 352
Subgeneric Names 356
Tribal and Ordinal Names 357
Names of Cohorts, Classes, &c 358
SECTION II. GLOSSOLOGY OR TERMINOLOGY 359
SECTION III. DESCRIPTION 861
Characters 361
Punctuation 364
Synonomy 365
Iconography 366
Habitat and Station, &c 366
Etymology of Names 366
Accentuation, Abbreviations 367
Signs 368
Floras, Monographs, &c 369
SECTION IV. SPECIMENS, DIRECTIONS FOR THEIR EXAMINA-
TION, PRESERVATION, &c 370
Implements of Investigation 370
Diagrams 371
Herborizing 371
Drying Specimens 375
Poisoning Specimens 379
The Herbarium 380
ABBREVIATIONS 385
SIGNS 391
GLOSSARY OF BOTANICAL TERMS, WITH INDEX . . 393
STRUCTUBAL BOTANY
THE BASIS OF MORPHOLOGY.
INTRODUCTION.
1. THE two Biological Sciences, 1 considered as parts of Natural
History, are Zoology and Botany. The latter is the natural
history of the Vegetable Kingdom. It embraces every scientific
inquiry that can be made respecting plants, their nature, their
kinds, the laws which govern them, and the part they play in
the general economy of the world.
2. We cannot distinguish the vegetable from the animal king-
dom by any complete and precise definition. Although ordinary
observation of their usual representatives may discern little that
is common to the two, yet there are many simple forms of life
which hardly rise high enough in the scale of being to rank dis-
tinctively either as plant or animal ; there are undoubted plants
possessing faculties which are generally deemed characteristic of
animals ; and some plants of the highest grade share in these
endowments. But in general there is a marked contrast between
animal and vegetable life, and in the part which animals and
plants respectively play in nature.
3. Plants only are nourished upon mineral matter, upon earth
and air. It is their peculiar office to appropriate mineral mate-
rials and to organize them into a structure in which life is mani-
fested, into a structure which is therefore called organic. So
the material fitted for such structure, and of which the bodies
1 Biology, the science of life, or rather of living things, in its earlier use
was equivalent to physiology : recently, it has come to denote the natural
history of plants and animals, t. e. of the two organic kingdoms, including
both their physiology and descriptive natural history.
1
2 INTRODUCTION.
of plants and animals are composed, is called organic matter.
Animals appropriate and live upon this, but have not the power
of producing it. So the vegetable kingdom stands between the
mineral and the animal ; and its function is to convert materials
of the one into food for the other. Although plants alone are
capable of building up living structure out of mineral mate-
rials, and are the sole producers of the organic matter which
is essential to animal life, and although animals consume that
which plants produce, yet plants also consume organic matter,
more or less, acting in this respect like animals in all their opera-
tions, except in the grand and peculiar one by which they
assimilate mineral matter. Most plants of the higher grades
assimilate largely and consume little, except in special opera-
tions. Some, on the contrar} r , are mainly consumers, and feed
upon formed organic matter, living in this respect after the
manner of animals. The living substance of plants and animals
is essentially the same.
4. Botany deals with plants : 1. As individuals, and in respect
to their structure and functions. 2. In their kinds, and as
respects their classification, nomenclature, &c. Accordingly,
the most comprehensive division of the science is into PHYSIO-
LOGICAL or BIOLOGICAL BOTANY (using these terms in their widest
sense) and SYSTEMATIC BOTANY. But as Plrysiolog}' and Biology,
in the restricted sense, relate only to functions or actions and
their consequences, the first department naturally divides into
two, viz. Structural Botany and Physiology.
o. STRUCTURAL BOTANY comprehends all inquiries into the
structure, the parts, and the organic composition of vegetables.
This is termed ORGANOGRAPHY, when it considers the organs or
obvious parts of which plants are made up, and MORPHOLOGY,
when the study proceeds on the idea of type. The term
ORGANOGENY has been applied to the study of the nascent
organs and their development; PHYTOTOMY, or VEGETABLE
ANATOMY, to that of the minute structure of vegetables as re-
vealed by the microscope, i. e. to the composition of the organs
themselves. But, since anatomy in the animal kingdom includes
the consideration of general as well as of minute structure, and
indeed answers to organograplrv, the minute anatomy of both
kingdoms takes the special name of HISTOLOGY. The study of
functions, or of the living being (animal or plant) in action,
is the province of PHYSIOLOGY.
G. SYSTEMATIC BOTANY, or the study of plants in their kinds
and in regard to their relationships, comprises TAXONOMY, or the
principles of classification, as derived from the facts and ideas
INTRODUCTION. S
upon which species, genera, &c., rest; CLASSIFICATION or the
SYSTEM OF PLANTS, the actual arrangement of known plants in
systematic order according to their relationships ; PHYTOGRAPHY,
the rules and methods of describing plants ; and NOMENCLATURE,
the methods and rules adopted for the formation of botanical
names. GLOSSOLOGY or TERMINOLOGY 1 is a necessaiy part of
Phytography or Descriptive Botany, and hardly less so of
Structural Botany: it relates to the application of distinctive
terms or names to the several organs or parts of plants, and to
their numberless modifications of form, &c. This requires a
copious vocabulary of well-defined technical terms, b}' the use of
which the botanist is able to describe the objects of his study
with a precision and brevity not otherwise attainable. It will
be convenient to exemplify the principal terms along with the
modifications of conformation which they designate ; and also,
for greater fulness and facility of reference, to append to this
volume an alphabetical summary of them, or Vocabulary of
Botanical Term s . 2
7. The present volume is mainly devoted to Morphological
Botany ; that is, to Structural Botany on the basis of mor-
phology. This department cannot be properly dealt with apart
from considerable reference to intimate structure, development,
and function, the subject-matter of vegetable histology and
physiology. But these will here be treated only in the most
general or incidental and elementary way, and only so far as
is necessary to the understanding of the morphology of the
stem, leaves, &c. The whole discussion of the histology and
physiology of plants is relegated to a following volume and to
another hand.
8. The most comprehensive and important division of the
vegetable kingdom is into plants of the higher and of the lower
series or grade, i.e. into PH^NOGAMOUS (or PHANEROGAMOUS) or
FLOWERING, and CRYPTOGAMOUS or FLOWERLESS PLANTS. The
first are all manifestly of one type, and therefore have a consist-
ent and simple morphology. The second differ among them-
selves almost as widely as they do from the higher series ; and
1 GLOSSOLOGY is the better word, but TERMINOLOGY, although a hybrid
of Latin and Greek, is in common use.
2 What is called GEOGRAPHICAL BOTANY is the study of plants in respect
to their natural distribution at the present time over the earth's surface, and
the causes of it. FOSSIL BOTANY (Vegetable Palaeontology) relates to the
plants of former ages, as more or less made known in their fossil remains.
MEDICAL BOTANY, AGRICULTURAL BOTANY, and the like, are applications
of Botany to medicine, agriculture, &c.
4 INTRODUCTION.
their morphology is more special and difficult. Wherefore it is
better to treat them separately and subsequently. This will be
done in a third part, by an associate devoted to Cryptogamic
Botany.
9. Thus the field is here left clear for the Structural Botany
of Phaenogarnous or Flowering Plants, with which the study of
the science should naturally begin. In theory it may seem
proper to commence with the simplest plants and the most ele-
mentar}- structures ; but that is to put the difficult and recondite
before the plain and obvious. The t}*pe or plan of the vegetable
kingdom, upon which morphological botany is grounded, is fully
exemplified only in the higher grade of plants, is manifest to
simple observation, and should be clearly apprehended at the
outset.
CHAPTER I.
OUTLINES OF THE GENERAL MORPHOLOGY OF
PH^ENOGAMOUS PLANTS.
10. MORPHOLOGY, the doctrine of forms, as the name denotes,
is used in natural history in nearly the same sense as the older
term Comparative Anatomy. If it were concerned merely with
the description and classification of shapes and modifications,
it would amount to little more than glossology and organography.
But it deals with these from a peculiar pmnt of view, and under
the idea of unity of plan or type. 1
11. As all vertebrate animals are constructed upon one type
(or ground plan) , which culminates or has its archetype in man,
so all plants of the higher grade (8) are strictly of one type ;
the different kinds being patterns or repetitions of it, with varia-
tions. The vegetable kingdom, however, does not culminate in
an archet} T pe or highest representative. As respects the organs
of vegetation, the higher classes of cryptogamous plants exhibit
this same type ; but it is only in the most general or in a
recondite sense that this can be said of their organs of repro-
duction, and of the less differentiated structure of the lowest
classes. Wherefore ciyptogamous plants are left out of the
present view, to be treated apart.
12. Viewed morphologically and as to its component organs,
a plant is seen to consist of an axis or stem, which sends off
roots into the soil, and bears lateral appendages, commonly as
leaves, but which may be very unlike leaves in whole appearance
1 The term Morphology was introduced into science by Goethe, at least as
early as the year 1817 (Zur Naturwissenschaft uberhaupt, besonders zur
Morphologic, Stuttgart und Tubingen, 1817-24). On page 9 of the first
volume, he is understood to have suggested this word for the purpose and in
the sense now adopted in botany and zoology. It essentially replaces an
earlier and somewhat misleading word, Metamorphosis. (304.)
Apparently the first botanist to adopt the term was Auguste de St.
Hilaire, in his " Lemons de Botanique, comprenant principalement la Mor-
phologie Vegetale, etc., Paris, 1841. The term seems not to have been taken
up, in zoology, by Etienne Geoffrey Saint-Hilaire, the antagonist of Cuvier
(who was of a wholly different family from that of the botanist), although
the same idea was denoted by his phrase " unity of organic composition "
6 GENEKAL MOKPHOLOGY
and function. These appendages, whatever their form or use,
accord with leaves in mode of origin, position, and arrangement
on the axis or stem. Their most general and ordinary form is
the familiar one of foliage ; hence the name of leaves has been
by botanists extended in a generic way from the green expan-
sions which constitute foliage to other forms under which such
appendages occur. The proper morphological expression is,
that the latter are homologous with leaves, or are the homologues
of leaves. 1
13. Leaves are borne upon the stem at definite places, which
are termed NODES. A node may bear a single leaf or a greater
number. When it bears two, they occupy opposite sides of
the stem. When three, four, or more, they divide the circum-
ference of the stem equally, forming a circle, technically a
WHORL, or in Latin form a VERTICIL. When only two, the pair
evidently answers to the simplest kind of whorl. So that leaves
are either single on the nodes, in which case they are alter-
nate, that is, come one after another on the stem ; or in whorls
(whorled, verticillate) , in the commoner case of a single pah-
being called opposite. The bare space between two successive
nodes is an INTERNODE. This is longer or shorter, according to
the amount of longitudinal growth, which thus spaces the leaves,
or whorls of leaves, in most .various degrees, either widely when
the internodes are elongated, or slightly when they remain very
short. The plant, therefore (roots excepted) , is made up of a
series of similar parts, i. e. of portions of stem, definitely bearing
leaves, each portion developed from the apex of the preceding
one. This constitutes a simple-stemmed plant.
14. Branching is the production of new stems from the older
or parent stem. These normally appear in the AXILS of leaves,
that is, in the upper angle which the leaf forms with the stem,
from which they grow much as the primary stem grew from the
seed. The primary stem, connected with the ground, produces
roots which develop downwardly into the soil, from which they
draw sustenance. Branches, when developed above ground,
1 A common designation for all these appendages being desirable, a good
one is furnished by the Greek name for leaf, tyvXXov, PHYLLI M, plural
PHYLLA. This, used with prefixes, may be made to designate the kind of
leaves in many cases, as, prophylla, cataphi/lla, hgpxojtkylla.
Recent German botanists use the word Phyllome. in this sense. It is a
rather convenient and well sounding word ; but pkylloma is the exact Greek
equivalent of our word foliage, and therefore not very well chosen as a
common term for leaves which are not foliage as well as those which are.
Nor will this word, like phi/Hum, readily take prefixes, as above, or the adjec-
tive form, as it readily does in prophyllous, hyi>sophyllous, cjamophyllous, e.
OF PH^ENOGAMOUS PLANTS.
being in organic connection with their parent stem, do not
usually produce roots ; but when placed in equally favorable
conditions for it, i. e. on or in the soil, they may strike root as
freely as does the original stem.
15. An incipient stem or branch, with its rudimentary leaves,
is a BUD. The normal situation of a bud is in the axil of a leaf
(axillary) , the development giving rise to branches ; or else at
the apex of an axis (terminal), where there can be only one, the
development of which continues that axis. 1
16. As branches are repetitions and in one sense progeny of
the stem which bears them, so the serial similar parts or leaf-
bearing portions of a simple stem are repeti-
tions, or in a like sense progeity, each of the
preceding one from which it grew. The
simple-stemmed plant is made up of a series
of such growths, each from the summit of
its predecessor ; the branched plant, of ad-
ditional series, laterally developed, from ax-
illary buds. These ultimate similar parts into
which a plant may thus be analyzed, and
which are endowed with or may produce all
the fundamental organs of vegetation, were by
Gaudichaud called PHYTONS. But phyton,
being the common Greek name for plant, was
not a happily chosen appellation for plant-
elements, or homologous plant-units. A better
term for them is PHYTOMERA (qivrov, plant,
jM
n above is the caulicle.
12
MORPHOLOGY
After this is consumed and in good part converted into struc-
ture, the plantlet must by the action of its root and leaves imbibe
from the soil and air appropriate materials, and
assimilate them into nourishing matter needful
for further growth. Only then does the rudi-
ment of new structure appear, in the form of a
growing point, or bud, at the node or apex of
the primitive stemlet, between the two seed-
leaves. In this case it soon shows itself as
a second pair of leaves, at first resting on
the node (Fig. 9), next as somewhat upraised
by the development of the second internode
(Fig. 10, summit), and finally both this inter-
node and the pair of leaves complete their
growth (Fig. 11). Then the terminal bud
which crowns the second node develops in
the same way the third pair of leaves and
their supporting internode or joint of stem
(Fig. 12) ; and so on.
24. The root and the stem grow not only
in opposite directions, but in a different mode.
The primordial stem, pre-existing in the seed
(though at first it may be extremely short)
grows throughout its whole length, but most
in its upper part, so that it may become a
stemlet two or three inches long. But, soon
attaining its full growth as to length, the
stem is carried upwards by the subsequent joints or portions,
similarly developed and elongated, one after the other. Not
that each portion necessarily waits until the growth of its prede-
cessor is complete, though this occurs at first in seedling Maples
and other embryos unprovided with much store of food, yet the
development follows this course and order of succession. The
root, on the contrary, cannot be said to pre-exist in the seed, or
at most it ma}~ be said to exist potentially in tissue of the caulicle
from which a root or roots normally originate. 1 It is formed
1 Yet from nothing which is special to this part of the embryo, nor to the
embryo at all. The primary root is developed from subjacent tissue of the
tip of the caulicle, just as it is sometimes developed from along the sides,
and as secondary roots are from all or most stems under favoring conditions.
This complete similarity, and the fact of what is called the " endogenous "
origin of roots (i,e. their springing from subjacent rather than superficial
tissue) appear fully to warrant the statement in the text above.
FIG. 10. Maple plantlet with second internode developing. 11. Same with second,
internode an:l rxiir of leaves co:in5ete, and bud of the third apparent.
OF THE EMBRYO AND SEEDLING.
13
in the process of germination, and originates in tissue just back
of that which covers the root-end of the caulicle, and which,
being carried forward by the subjacent
formation (to which it becomes a sort of
cap or sheath) , is called the Root-cap.
As the primary root thus began by a
new and local growth at the extremity
of pre-existing stem, so it goes on to
grow in length wholly or mainly by a
continuation of this formation, the new
at the end of the old. That is, the
root elongates by continual minute
increment of its apex or near it, the
formed parts very soon ceasing to
lengthen. This is in marked distinc-
tion from stem, which grows by suc-
cessive individualized portions ; and
these portions (internodes) , at first
very short, attain or are capable of
attaining a considerable and sometimes
very great, but definitely terminable
length, by interstitial growth through-
out. Moreover, roots are naked, not
producing as they grow either leaves
or any organs homologous with leaves.
The} T commonly branch or divide, but
in a vague manner ; and their new parts bear what are called
Root-hairs, which greatly increase the absorbing surface ; other-
wise they are destitute of appendages or organs.
25. With the Maple embryo, here taken as a type, that of
Morning Glory, Ipomcea purpurea, or any of its kin, may next
be compared. The cot}'ledons are different in shape, being as
broad as long, and notched both at base and apex. They lie
in contact in Fig. 14, and are very thin, leaf-like, and green
while contained in the seed. Their thinness is shown in Fig. 13,
where a section of the crumpled and folded embryo, as it lies in
the seed, exactly divides them (passing through the terminal and
basal notches) and also the caulicle, which here is thicker than
both. The germination is similar to that of the Maple ; and like
that (as Fig. 16 shows), and for the same reason, no bud or
rudiment of the further growth pre-exists in the embryo or
FIG. 12. Red Maple seedling, with throe joints of stem and pairs of leaves developed,
the first being the cotyledons.
MORPHOLOGY
appears in the young plantlet, until that has established itself
and had time to elaborate proper material therefor. This con-
dition is correlated with thin foliaceous
cotyledons, holding no store of nourish-
ment. Here they do not contain sufficient
material for the development of the initial
stem and root. The maternal provision
for this is here stored up in the seed
around but not within the embryo. This
nourishing deposit, seen in the section
(Fig. 13) filling the whole space between
the seed-coats and the thin embryo, was
named by the early botanists and vege-
table anatomists the ALBUMEN of the seed. 1
This substance, softened in germination
and by chemical changes rendered soluble,
is gradually absorbed by the cotyledons
as material for their growth and that of
the developing primary stem and root.
26. Seeds in this regard are accordingly
distinguished into albuminous and exal-
buminous, those supplied with and those
destitute of albumen. The difference
inheres neither in the character nor in
the amount of the maternal provision for the development of
the embryo-plant, but merely in the storage. In exalbuminous
seeds the nourishment supplied for this purpose is taken into
the embr} T o itself, mostly into the cotyledons, during the growth
and before the maturity of the seed. In albuminous seeds
this same material is deposited around or at least external to
the embryo.
27. The amount of this deposit is, in the main, inversely pro-
1 Grew appears to have first applied this name, and Gaertner to have
introduced it into systematic botany, where it remains in use, although
Jussieu replaced it by the term Perisperm, and Richard by Endosperm,
neither of them much better etymologically than the old word Albumen.
But it must be kept in mind that it was intended to liken the " albumen " of
the seed with the albumen or white of an egg as a body or mass, and not as
a chemical substance; the embryo being fancifully conceived to be analo-
gous to the yolk of the egg, the surrounding substance of this kind not
unnaturally took the name of the white, viz. albumen.
FIG. 13. Section of seed of common Morning Glory, Ipomoea purpurea, dividing
the contained embryo through the centre. 14. Embryo of same, detached and straight-
ened. 15. Embryo in germination ; the cotyledons only partly detached from the coat
of the seed. 16. Same, later and more developed, the cotyledons unfolded and out-
spread as the first pair of leaves.
OF THE EAIBKYO AND SEEDLING.
15
portional to the size and strength of the embiyo, or the degree
of its development in the seed. A comparison of the various
illustrations sufficiently shows this. Figures 17 to 24 exhibit,
in a few common
seeds, somewhat of
this relation, and
also of the position
and shape assumed
in some instances.
The tipper rank of
figures represents
sections of seeds ;
the embiyo left in
white ; the albumen
as a dotted surface.
The lower rank shows the embryos detached. That of Mirabilis
has very broad and thin cot}'ledons, a caulicle of equal length,
and the whole curved round the albumen which thus occupies
the centre of the seed. That of Potato is coiled in the midst
of the albumen, is slender ; the cotyledons narrowed down to
semi-cylindrical bodies, not leaf-like in appearance, and the two
together not thicker than the caulicle. In Barberiy the embiyo
is straight, in the axis of the albumen, which it almost equals in
length ; the cotyledons considerably broader than the caulicle,
but short and thickish. That of the Peony is similar, but very
much smaller, occupying a small space at one end of the albu-
men, and seemingly without distinction of parts, but under the
microscope and with some manipulation the broader end is
found to be divided, that is, to consist of two minute cotyledons.
The embiyo of a Crowfoot is similar, but still more minute and
the parts hardly to be distinguished ; and in some minute em-
bryos there is no apparent distinction of parts until they develop
in germination.
28. The study of the formation of the embryo in the seed
teaches that all embryos begin with a still more simple, minute,
and homogeneous structure ; and these comparisons suffice to
show that all such differences are referable to different degrees
and somewhat different modes of the development of the embiyo
while }-et in the seed. It also appears that the size and shape
FIG. 17. Section of seed and contained embryo of Mirabilis (Four-o-clock).
18. Embryo detached entire.
FIG. 19. Section of a Potato-seed. 20. Embryo detached entire.
FIG. 21. Section of Barberry-seed. 22. Embryo det.ac.hed entire.
FIG. 23. Section of Peony-seed. 24. Embryo detached entire.
16
MOKPHOLOGY
of an organ do not indicate its nature, either in the embn^o or
in subsequent growth. But in all the cases } T et mentioned the
cotyledons actually demonstrate their
nature by developing in germination
in a foliaceous manner and becoming
the first leaves of the seedling. Nor
is this nature much disguised by the
fact that they differ greatly in form in
different species, and that the seed-
leaves, or developed cotyledons, differ
much in shape and often in texture
from the succeeding leaves. (See Fig.
11, 12, 25, &c.)
29. To complete the comparison
between the seedling Morning Glory
and that of the Maple, it is to be
noted that here, while the cotyledons
or seed-leaves are two, the following
internode bears only one leaf (Fig. 25), as also will the just de-
veloping third internode ; and this continues throughout up to
the blossom : that is, the leaves subsequent
to the cotyledons are not opposite as in the
Maple, but alternate. (13.)
30. All the preceding illustrations are from
embryos w r hich previous to germination have
developed nothing beyond the cotyledons. In
the following, a rudiment of further growth,
or a primary terminal bud, is visible in the seed. It is most
manifest in large and strong embryos with thick or fleshy cotyle-
FIG. 25. Further development of Morning Glory, Fig. 1G, the root cut away, the
internode above the cotyledons and its leaf completed, the next internode and its leaf
appearing.
FIG. 26. Embryo (kernel) of the Almond. 27. Same, with one cotyledon removed,
to show the plumule, a.
FIG. 23. Section of an Apple-seed, magnified, cutting through the thickness of the
cotyledons. i>9. Embryo of the same, extracted entire, the cotyledons a little separated.
FIG. 30. Germination of the Cherry, showing the thick cotyledons little altered,
and the plumule developing the earliest real foliage.
OF THE EMBRYO AND SEEDLING.
17
dons, i. e. cotyledons well charged with nourishing matter. The
early vegetable physiologists gave to it the name of PLUMULE
(Lat. plumula, a little plume). The K t
name was suggested by its appearance
in such an embr3'o as that of the bean
(Phaseolus), in which it evidently con-
sists of a rudimentary pair of leaves,
while in the pea and the acorn it is a
rudimentary stem, the leaves of which
appear only later, when germination
has considerably advanced. In any
case, the plumule is the bud of the
ascending axis already discernible in
the seed. Fig. 27, a, shows it in the
almond, one cotyledon being removed.
Fig. 28 shows it in the section of a
similar although much smaller embryo,
that of an apple-seed, enlarged to
nearly the size of the other. It is
equall}' visible in the cherry, the bean,
and the beechnut. The embryo in all
these cases constitutes the whole kernel
of the seed. For the nourishment,
which in all the foregoing illustrations
except the first (i.e. in Fig. 13, 17-23),
is deposited around or exterior to the
embiyo, is in these stored within it.
31. The development of these em-
bryos in germination proceeds in the
normal manner, but with two cor-
related peculiarities. First, b} r the
lengthening of the radicle more or less, their thick cotyledons
are usually raised to or above the surface of the soil ; they
expand, assume the green color needful to foliage ; but the}'
imperfectly or in a small degree perform the function of
green leaves. Their main office is to supply the other growing
parts with the prepared nourishment which they abundantly
contain. Then, being thus copiously nourished, the root below
and the read3'-formed plumule above grow rapidly and strongly,
having accumulated capital to draw upon ; and the leaves of the
FIG. 31. Beechnut cut across, filled by the fli shy embryo; the thick cotyledons
partly enfolding each other. 32. Embryo of the same in early germination 33. Same
more advanced; the plumule, which is just emerging in the preceding, here developed
into a long internode and a pair of leaves.
. 2
18
MORPHOLOGY
latter are practically the earliest efficient foliage of the plantlet.
Thus, as in the germinating Cheny-seed (Fig. 30) , three or four
internodes of stem, with their leaves, ma}^ be produced before
these leaves themselves are sufficiently developed to make any
sensible contribution to this growth. And in the Beech and Bean,
the leaves of the plumule come forward almost before the root
has attached the plantlet to the soil. (Fig. 32, 35.) Between
such cases and that of Maple and the like there are all degrees.
There are also familiar cases in which the storage of nourishment
in the cotyledons is carried to a maximum, with results which
gravely affect the development.
FIG. 34. The embryo (the whole kernel) of the Bean. 35. Same early in germi-
nation ; the thick cotyledons expanding and showing the plumule. 36. Same, more
advanced in germination ; the plumule developed into an internode of stem bearing a
pair of leaves.
FIG. 37. Embryo of Pea, i. e. a pea minus the seed-coat. 38. Advanced germi-
natior of the same.
OF THE EMI3KYO AND SEEDLING.
19
32. Thus, in the Pea, near relative of the Bean, the embryo
(Fig. 37), which is the whole kernel of the seed, has the
cotyledons so gorged with this nutritive
store that they are hemispherical ; and
the acorn of the Oak (Fig. 39), near
relative of the Beech, is in similar case.
These extremely obese cotyledons have
not only lost all likeness to leaves, but all
power of fulfilling the office of foliage,
which is apparently no disadvantage ; for
when two different duties are performed
by the same organ, it rarely performs both
equally well. Here they become mere
receptacles of prepared food, the nature
and office of which is the same as of the
albumen, or nutritive deposit exterior to
the embryo in what are called albuminous
seeds. (25-27.) The difference is in the
place rather than in the character of the
deposit. The plumule in such cases is
always apparent before germination ; and
it develops even with more vigor than in
the preceding cases. It usually rises as a
stout stem of several internodes lengthen-
ing almost simultaneously, or at least the
upper strongly developing long before the
lower have finished their growth ; and
the latter are practically leafless, bearing
only small and scale-like and useless ru-
diments of leaves. This is correlated with
the peculiarity that the caulicle does not
lengthen in germination, or it lengthens
very slightly ; the cotyledons remain within
the coats of the seed ; and if this were ^
buried beneath the surface of the ground, there it remains. The
abortion of the earliest leaves of the plumule is in correlation
with this hypogceous (i. e. underground) situation of the cotyle-
dons throughout the germination. The slight elongation of the
caulicle S3rves merely to protrude its root-end from the coats of
the seed in a downward direction, and from this a strong root
usually is formed.
FIG. 39. Section of an acorn, filled by the embryo. 40. Advanced germination oi
the same.
20 MOIU'HOLOGY
33. In some Oaks, notably in our Live Oak (Quercus virens),
and less so in the Horsechestnut, the two cotyledons coalesce or
cohere by their contiguous faces.
In some of these cases of hypo-
gaeous germination, the short
caulicle and plumule are extri-
cated from the enclosing coats or
husk by the development of short
stalks (petioles, 157) to the fleshy
cotyledons ; as is seen in Fig. 42,
and in most germinating acorns.
These petioles are not visible in
the seed, but are the first develop-
ment in germination.
34. There are some curious
cases in which, while the caulicle
remains short and subterranean,
the cotjledons are raised out of
ground in germination by the
formation of far longer stalks
(petioles) than those of the
Horsechestnut. A singularl}* dis-
guised instance of this kind is seen in Megarrhiza, a genus of
Cucurbitaceous plants of California and Oregon, remarkable for
their huge root. The large seed has very thick and fleshy
cotyledons, and a veiy short and straight caulicle. In germi-
nation, the whole seed is elevated, seemingly in the manner of
tli2 bean, upon a stout stem. One waits for a long time expect-
ing to see the cotyledons throw off the bursting husk and expand,
or else to put forth the plumule from between their bases. But
at length the plumule makes its appearance from an unexpected
place, coining separately out of the soil. Removing this, the
state of things represented in Fig. 43 is presented, that of
the plumule seemingly originating from the base, instead of the
apex, of an elongated caulicle ! But on examination of the cleft
from which this proceeds, by making a section of the stem above
(showing that it is hollow) , and finally by separating the cotyle-
dons and gently tearing apart the two short stalks by which the}'
are united to their stem-like support, it is found that the latter may
be divided into two (as shown in Fig. 44) , even down to the cleft
below. This explains the anomaly. The real caulicle has re-
FIG. 41 Section of a Horsechestnut or Buckeye seed, through the very thick
cotyledons and the incurved caulicle 42. Seed in germination, showing the petioles
to the cotyledons, &c.
OF THE EMBRYO AND SEEDLING.
21
mained short and subterranean, and is confluent with the upper
part of the thickening root : the seeming caulicle, which raised
the cot} T ledons above the soil, consists
of the petioles of these combined into
a tubular stem-like body, no evident
trace of which is visible in the seed,
although in germination it attains the
length of two or three inches : in age
it is readily separable into the two
leaf-stalks or petioles of which it is
composed : the plumule is thus seen
to be wholly normal, originating from
between the cotyledons. All the ex-
tensive growth so far, and until the
proper foliage-leaves of the continu-
ation of the plumule are developed
and begin their action, is from nutri-
tive material stored in the thickened
cotyledons, a considerable part of
which was transferred to the already
enlarging root, before a remaining
portion was used in building up the
strong plumule. The economy of this
elevation of cotyle-
dons which never
open, and of the
lengthened distance
through which the
nutritive matter has
to be carried, is not
apparent. But it is
the family habit in
Cucurbitacese to
bring up the cotyle-
dons that they ma} r
develop as leaves
(as in the Pumpkin,
Fig. 47) : here this
elevation is brought
about in a different waj', but without securing the useful end. 1
1 It may be inferred that Megarrhiza is a descendant of some Cucurbitacea
with thinner cotyledons, which in germination developed into long-stalked
leaves, in the manner described in the next following paragraphs.
FIG. 43, 44. Peculiar germination of Megarrliiza California! ; explained above.
22
MORPHOLOGY
35. This same anomaly, as to the development of long stalks
to the cotyledons and their union into a stem-like bod}', occurs in
various species of Larkspur (notably in the Californian Delphin-
ium nudicaule) ; but in these the cotyledons develop into a pair
of efficient green leaves.
45 47
36. A similar elongation of petioles of the cotyledons, but
without any union, occurs in a species of Morning Glory of the
plains beyond the Mississippi (Ipomcea leptophylla) ; the leaf-
like cotyledons coming up on their long stalks separately from
the ground (Fig. 45) ; the developed plumule rising some
time afterward between them. Compare this with the ordinary
species (25, Fig. 15, 16, 25), and note that the difference is merely
that the caulicle in the common Morning Glory elongates and the
petioles of the cotyledons remain short.
37. In all instances thus far a single primar} r root so regularly
develops from the lower end of the axis of the embryo (variously
named radicle or caulicle) , and forms such a direct downward
FIG. 45. Germination of Ipomoea leptophylla; the caulicle not developing, the
plumule and the petioled cotyledons rise from underground. Dotted line marks the
level of the soil.
FIG. 46. Embryo of a Pumpkin, the cotyledons separated. 47. Same germinated;
a cluster of roots from the base of caulicle.
OF THE EMBRYO AND SEEDLING. 23
prolongation of it, that it was called the descending axis ; and
the body from which it originates was named the radicle, on the
supposition that it was itself the nascent root. But, as already
explained, the so-called radicle grows in the manner of stem (24),
and is morphologically that initial internode the node of which
Dears the first leaves or cotyledons. (20.) Let it now be noted
that this descending axis or single primary root is far from
universal. In Pumpkin, Squash, Echinoc}'stis, and the like,
the strong caulicle sends out directly from its root-end a cluster
of roots or rootlets, of equal strength; i. e., it strikes root in
nearly the manner that a cutting does. (Fig. 47.)
38. The Polycotyledonous Embryo is one having a whorl of
more than two seed-leaves. The dicotyledonous embryo being
a whorl of the very simplest kind, that is, with the
members reduced to two, the poly cotyledo nous
may be regarded as a variation of it. In all but
one group of plants it is simply a variation, of
casual occurrence, or even a monstrosity, in which
three or rarely four cotyledons appear instead of
two. In Pines (Fig. 48, 49), however, and in
most but not all Coniferse, a whorl of from 3 to 10
cotyledons is the normal structure, va^-ing accord-
ing to the species, but of almost uniform number
in each. In germination these are brought out of
the soil by the elongation of the caulicle, and when
the husk of the seed is thrown off they expand
into a circle of needle-shaped leaves. In the Pine
tribe, all the subsequent leaves are alternate (spiral) in arrange-
ment, with some disguises. In the C}'press tribe, the cot3 T ledons
are fewer (not more than four, and more commonly only two),
and the subsequent leaves also are in whorls of two to four ;
i. e., are either opposite or verticillate. From the occasional
union at base of the cotyledons of a polycotyledonous embryo in
pairs or groups, and from a study of their earl} 7 development,
Duchartre 1 plausibly maintains that such cotyledons realty consist
of a single pair, parted into divisions or lobes. The ordinar}'
interpretation, however, is equally tenable.
39. The Monocotyledonous Embryo, although theoretically the
simplest, is practically a more difficult stud}'. It has a single
cotyledon (as the name denotes) ; also a single leaf to each node
1 Ann. Sci. Nat. ser. 3, x. 207. This view, which originated with Jussieu,
is adopted by Parlatore in DC. Prodr. xvi.
FIG. 48. Section of a seed of a Pine, with its embryo of several cotyledons. 49. Early
seedling Pine, with its stemlet, displaying its six seed-leaves.
MORPHOLOGY
64
of the plumule ; that is, the leaves of the embryo are alternate.
But the caulicle is usually very short, and there is no external
mark by which its limits may be distin-
guished from the cotyledon, until germi-
nation has begun. For a type of it, the
embryo of some aquatic or marsh plants
\i \ // |Nj2>d| may be taken, where it forms the whole
kernel of the seed (Fig. 50-53), and
50 si 52 53 the structure can be made out antecedent
to germination. It is understood by supposing that the cotyle-
don, which forms its principal bulk (the caulicle being only the
very short thickish base) , is convolute around a short
plumule, and the margins concreted, except a minute
longitudinal chink at base, out of which the growing
plumule protrudes in germination. The embryo of
Iris may be similar in structure, but no distinction
of parts is visible. It is very small in proportion to
the size of the seed, the kernel being mostly albu-
men, a supply of food, from which the germinating
embryo draws the materials of its growth. When
this takes place, either the cotyledon or the whole
embryo lengthens, its lower part is pushed out of the
seed, a root forms at the free end of the excessively
short caulicle, and the plumule develops from the
other in a series of one-leaved nodes, the internodes
of which remain so short that the leaves continue
in close contact, the bases of the older successively
enclosing the inner and younger. (Fig. 55.) Here,
therefore, the cotyledon mainly remains in the seed,
and the seed remains underground (Irypogseous) .
40. It is somewhat different in the Onion, which
has a similar embryo, except that it is longer, and
the cotyledon is curved in the albumen of the seed.
The first steps are the same as in Iris ; but as soon
as a root is formed and embedded in the soil, the
cotyledon lengthens vastly more, into a long and
filiform green leaf, which, taking an erect position,
FIG. 50. Seed of Triglochin palustre ; the rhaplie, leading to the strong chalaza at the
summit, turned towards the eye 51. The embryo detached from the seed-coats, showing
the longitudinal chink at the base of the cotyledon ; the short part below is the radicle.
52. Same, with the chink turned laterally, and half the cotyledon cut away, bringing t o
view the plumule concealed within. 53. A cross-section through the plumule, more
magnified.
FIG.' 51 Section of seed of Iris, enlarged, showing the small and apparently simple
embryo at the base of the albumen 55 Germinating seed and seedling of the same, of
natural size.
OF THE EMBRYO A!ND SEEDLLNG.
25
carries up the light seed far above the surface of the ground, the
tip only remaining in the albumen of the seed until that is ex-
hausted, when the tip perishes and the emptied husk falls away.
About this time the plumule shoots forth from one side of the
subterranean base of this cotyledonar leaf, in the form of a second
and similar filiform leaf, to be followed by a third, and so on.
The sheathing bases of these succeeding leaves become the coats
of the Onion-bulb. The internodes remain undeveloped until the
plant is ready to blossom. Very similar is the germination of
a date-seed, except that the
protruding cotyledon does
not lengthen so much, nor
does it elevate the heavy
seed. Instead of the seed
being carried up, the lower
end of the embiyo, contain-
ing the plumule, is pushed down more or less into the loose
soil, from which in time the developing plumule emerges.
41. The embryo of Grasses, especially of those which yield
the cereal grains, is more complex, owing mainly to the great de-
velopment of the plumule
and the manner in which
its rudimentary
successively enclose each
other. That of Maize or
Indian Corn, one of the
largest, is most convenient
for study. (Fig. 56-59.) The floury part of the seed, which
makes most of its bulk, is the albumen, largely composed of
starch. The embryo is exterior to this, applied to one of its
flat sides, and reaching from the thinner edge to or above the
middle in the common variety of corn here represented. The
form of the embiyo is best shown, detached entire, in Fig. 58 :
its structure appears in the sections. The outer part is the
cotyledon, which incompletely enwraps the plumule : it adheres
closely to the albumen b}' the whole back, and remains un-
changed in germination : its function is to absorb nutritive
FIG. 56. Section, flatwise, of a grain of Indian Corn, dividing the albumen and the
embryo. 57. Similar soction at right angles to the first. 58. A detached embryo:
corresponding parts of Fig. 57 and 58 indicated by dotted lines.
FIG. 59. Vertical section of Indian Corn across the thickness of the grain, dividing
the embryo through the centre and displaying its parts: c, cotyledon; p, plumule;
r, the radicle or caulicle.
FIG. 60. Similar section of grain of rice. 61. Same of an oat-grain; the parts
as in Fig. 59.
26
MORPHOLOGY
matter furnished by the albumen, and to transmit it to the
growing plumule. The plumule consists of a succession of
rudimentary leaves, sheathing
and enclosing one another, on
the summit of a very short
axis, which is mainry the
caulicle, otherwise called rad-
icle. This is completely en-
closed b}' a basal portion of
the cotyledon and of the
outermost leaf of the plu-
mule, which form a peculiar
sheath for it, named the
Coleorhiza* i. e. root-sheath :
consequently the first root or
roots have to break through
this covering. As in the Oak
and Pea (32), the very first or
outermost leaves of the plu-
mule develop imperfectly and not into
efficient foliage. The one in Fig. 62,
which encloses the rest in the early
growth, is left behind as a mere sheath
to the base of the following and more
perfect leaves : it is the same as the
lowest in Fig. 63. The leaves are first
developed : the internodes lengthen later, and the lowest lengthen
very little. Not rarely the first root starts singly from the tip of
the caulicle (Fig. 62, just as in Fig. 55) ; but others of equal
strength follow from any part of the caulicle, and soon from
the nodes above ; and no tap-root is ever formed.
42. A Pseudo-monocotyledonous embryo occasionally occurs ;
that is, one of the dicotyledonous t}'pe, of which one cotyledon is
wanting through abortion. This occurs in Abronia, a genus
related to Mirabilis, and bearing an embiyo very similar to that
represented in Fig. 17, 18, except that one cot}'ledon is absent.
The anomaly of an acotyledonous embryo occurs in Dodder, a
plant of the dicotyledonous type, but with both cotyledons
1 This, the Cokorhize of Mirbel, should not be confounded (as by some it
has been) with the "root-cap," or tissue which ordinary roots (whether
primary or secondary) break through in their development or carry on
their apex.
FIG. 62. Early germination of Indian Corn. 63. More advanced germination ot
same : roots produced from portion of stem above the cotyledon as well as below.
OF THE ORGANS OF VEGETATION. 27
actually wanting, a correlation with its parasitic mode of life.
(64, Fig. 78.)
43. The dicotyledonous and the monocotyledonous character
of the embryo is correlated with profound differences in the whole
ulterior development, as revealed in the structure of the stem,
leaves, and flower ; which differences mark the two great divisions
of Phasnogamous plants, viz. DICOTYLEDONES or DICOTYLEDONOUS
PLANTS, and MONOCOTYLEDONES or MONOCOTYLEDONOUS PLANTS,
names introduced into classification by Ray, and adopted by
A. L. Jussieu, in his Genera Plantarum.
CHAPTER III.
MORPHOLOGY AND STRUCTURE OF THE ORGANS OF THE
PLANT IN VEGETATION.
SECTION I. OF THE ROOT.
44. The Root, which has been called the descending axis, is
that portion of the body of the plant which grows downward,
ordinarily fixing the vegetable to the soil, and absorbing from it
materials which the plant ma}- elaborate into nourishment. As
already stated (24), the root grows in length by continuous
additions of new fabric to its lower extremit}', elongating from
that part only or chiefly ; so that the tip of
a growing root always consists of the most
newly formed and active tissue. It normally
begins, in germination, at the root-end of the
caulicle, or so called radicle. But roots soon
proceed, or may proceed, from other parts of
the stem, when this is favorably situated for
their production. The root does not grow
from its naked apex, but from a stratum
immediately behind it : consequently its blunt
or obtusely conical advancing tip consists of older, firmer, and
in part effete tissue. The tip of all secondary roots and rootlets
FIG. 64. Magnified tip of root of a seedling Maple (such as in Fig. 9), sufficiency
enlarged to indicate the cellular structure: a. the portion where growth is taking
place; b the older and firmer tip.
MORPHOLOGY OF THE ROOT.
is similarly capped or protected. 1 But the so-called root-cap is
seldom so distinct or separable as to deserve a particular name.
45. Nature of Growth, Cells. The development and growth of
the root, as of other organs, results from the development,
growth, and increase in number of certain minute parts, of which
the plant is built up. These component parts are so much alike,
at least in an earl} 7 stage, and are so obviously formed all on
one type, that they take one common name, that of CELLS.
These are the histological elements of plants, i. e. the units of
minute anatomical structure. While, in the morphology of the,
plant's obvious organs, analysis brings us to the phytomer (16)
as the individual element which by a kind of propagation
produces its like in a second phytomer, remaining however in
connection with the first, thus building up the general structure,
so, in an analogous wa}', each of the obvious parts each stalk
or blade or rootlet is microscopically determined to be com-
posed of these ultimate organic units, generally called cells.
The cell (cellula, by the French
conveniently termed cellule) is the
living vegetable unit, in the same
sense that the brick is the unit
of a brick edifice. To make this
analogy fairly complete, the
bricks should be imagined to
have a firm exterior or shell,
and a soft or at length hollow
- interior, also to be living when
incorporated into the structure,
and finally to be produced in the
forming structure by a kind of
propagation. The production or
increase in number of these cells by development from previous
ones, and their successive increase in size up to maturit}', arc
what constitutes vegetable growth. 12 The inspection through a
1 The notion that the tip of the root consists of delicate forming or
newly formed tissue, or bears some organ or structure of this nature (a
" Spongiole"), has hardly yet been eliminated from the text-books and popular
writings. It had no proper foundation in fact.
In Lc.mna, and in some other aquatics, and also in some aerial roots, this
older tissue often separates into a real root-cap, free at base, like an inverted
calyptra.
2 This, as to the structure, is the subject of Ilistohjy ; as to processes or
actions, the subject of Physiology; both to be treated in a separate volume.
FIG. 65, 66. Portions of surface of Fig. 64, more magnified, clearly displaying the
superficial cellular structure and the long processes from some of the cells, called root-
hairs, which abound on the upper part of Fig. 64.
MORPHOLOGY OF THE ROOT. 29
simple microscope of a slender young root, and of thin slices
of it immersed in water, ma}' serve to give a general though
crude idea of the vegetable cellular structure, sufficient for the
present purpose. Roots are naked ; that is, the}' bear no other
organs. When they send off branches, these originate from the
main root just as roots originate from the stem ; and in both
cases without much predetermined order. The ultimate and
very slender branches are sometimes called root-fibrils ; but
these are only delicate ramifications of the root. Like any
other part of the plant, however, roots ma}' produce hairs or
such like growths from the surface, which are wholly distinct
from branches. (383.)
46. Root-hairs. Roots absorb water, &c., from the soil by
imbibition through the surface ; that is, through the walls of the
cells, which are in a certain sense permeable to fluids, more readily
when young and tender, less so when older and firmer. Roots,
therefore, absorb most by their fresh tips and adjacent parts ;
and these are continually renewed in growth and extended fur-
ther into the soil. As the active surface of a plant above ground
is enormously increased by the spread of foliage, so in a less
degree is the absorbing surface of young roots increased by the
production of root-hairs. (Fig. 64, upper part, and more magni-
fied in Fig. 65, 66.) These are attenuated outgrowths of some
part of the superficial cells into capillary tubes (only one from
each cell) , closed at the tip, but the calibre at base continuous
with the cavity of the cell ; into which, therefore, whatever is
imbibed through the thin wall may freely pass. These appear
(as Fig. 64 shows) at a certain distance behind the root-tip.
Further back the older or effete root-hairs die away as the cells
which bear them thicken into a firmer epidermis.
47. To the general statement that roots give birth to no other
organs, there is this abnormal, but b}' no means unusual excep-
tion, that of producing buds, and therefore of sending up leaf}'
branches. Although not naturally furnished with buds in the
manner of the stem, yet many roots have the power of originat-
ing them under certain circumstances, and some produce them
habitually. Thus Apple-trees and Poplars send up shoots from
the ground, especially when the superficial roots arc wounded.
And the roots of Madura or Osage Orange so readily originate
buds that the tree is commonly propagated by root-cuttings.
48. Kinds of Roots. The root, commonly single, which origi-
nates from the embryo itself, is called the PRIMARY ROOT. (37.)
Roots which originate from other and later parts of the stem,
or elsewhere, are distinguished as SECONDARY ROOTS. But the
eO MORPHOLOGY OF THE ROOT.
latter are as normal as the primary root ; that is, to stems
so situated that the}' can produce them. Most creeping plants
emit them freely, usually from the nodes ; and so do most
branches, not too old, when bent to the ground and covered
with earth, thus securing the requisite moisture and darkness.
Separate pieces of young stems (cuttings) can commonly be
made to strike root. Upon this faculty of stems to originate
roots depends all propagation by division, Ivy laying or layering,
by cuttings, &c. It is mainly annuals and common trees that
naturally depend on the primaiy root ; and most of these can be
made to produce secondary roots. Even leaves and leaf-stalks
of some plants may be made to strike root and be used as
cuttings. (77.)
49. Duration. By differences in respect to this, cither the
root or the plant, as the case may be, is distinguished into
Annual, Biennial, or Perennial, according to whether life is contin-
ued for a single year or season, for two, or for a greater number.
The difference is not in all cases absolute or even well marked.
50. Annuals are plants which, springing from the seed, flower
and seed the same year or season, and die at or before its close.
They produce fibrous roots, either directly from the embryo and
succeeding joints of stem (as in Grasses, Fig. 63), or from a
persistent primary or tap-root, more or less thickened into a trunk
or divided into branches. The products of vegetation in all such
herbs are not stored in subterranean or other reservoirs, but arc
expended directly in new vegetative growth, in the production
of blossom, and finally in the maturation of fruit and seed.
This completed, the exhausted and not at all replenished indi-
vidual perishes.
51. But some annuals may have their existence prolonged by
not allowing them to blossom or seed. Others, with prostrate
stem or branches, may from these produce secondary roots,
which, forming new connections with the soil, enable the newer
growth to survive when the older parts with the original root
have perished. And man}' herbs, naturally annuals, are continued
from year to year through such propagation from the branches,
used as layers or cuttings. Moreover, certain plants (such as
Ricinus or Castor-oil Plant) , which arc perennial or even arbo-
rescent in warm climates to which they belong, become annuals
in temperate climates, early perishing by autumnal cold.
52. The annuals of cool climates, where growth completely
ceases in winter, germinate in spring, mature, and die in or before
autumn. But, in climates with comparatively warm and rainy
winter and rainless summer, many germinate in autumn, vegetate
MORPHOLOGY OF THE BOOT.
31
through the winter, flower and seed in spring, and perish in
early summer. These may be termed WINTER ANNUALS.
53. Biennials are plants which, springing from the seed and
vegetating in one season, live through the interruption of winter,
and blossom, fructify, and perish in the next growing season ;
their life being thus divided into two stages, the first of vegeta-
tion, the second of fructification. In typical biennials, nearly
the whole work of vegetation is accomplished in the first stage,
with the result of accumulation of a stock of nutritive matter,
to be expended in the second stage in the production of blossom
and seed. This accumulation is usually stored in the root or in
the base of a ver}^ short stem in c :>nnection with the root. The
root of a biennial accordingly enlarges and becomes fleshy, or
obese, as this matter accumulates. At the close of the growing
season, the leaves perishing and the
stem having remained very short (with
undeveloped intsrnodes) , the root,
crowned with the bud or buds, contains
the main result of the summer's work,
as provision for the next year's devel-
opment and the completion of the
C}*cle. This development, being thus
amply provided for, is undertaken in
spring with great vigor ; blossom, fruit,
and seed are rapidl}' produced ; and
the stock being consumed, but not at
all replenished, the cells of the great
root are now empty and effete, and
the individual perishes. The Beet,
Turnip, Parsnip, and Carrot arc fa-
miliar examples of biennials, with the
store of nourishment in the root. 1
The Kohl-rabi is a biennial with this deposit in the stem :
the Cabbage, partly in the stem, partly in the head of leaves.
1 In these the caulicle enlarges with the root, so that the upper and
bud-bearing end is stem.
Tap-roots of this kind are said, in descriptive botany, to be
Fusiform or Spindle-shaped, when broader in the middle and tapering
towards both ends, as in the common Radish (Fig. 67);
Conical, when tapering regularly from base to tip, as in carrots, &c. ;
Napiform, i. e. Tiir)iij)-shaped,\\'\-\en the thickened part is wider than high, &c.
Fascicled Roots are those which form in clusters ; these may be slender or .
thickened. When much thickened, either irregularly or not of the above
shapes, they are said to be tuberous.
FIG. G7. Radish : a fusiform tap-root.
MORPHOLOGY OF THE ROOT.
54. But some plants, such as the Radish, which when they
spring from seed in autumn are true biennials, will when raised
in spring pass on directly to the flowering stage in summer, or
when sown after the warm season begins will often run through
their course as annuals. Then there are various biennials which
thicken the root very little and hold their leaves through the
winter. Between these and winter annuals no clear demarcation
can be drawn. As respects annual and biennial duration, the
terms may for the most part be applied indiscriminately to the
plant or to the root. We may say either that the plant is a
biennial, or that its root is biennial.
55. Perennials are plants which live and blossom or fructify
year after year. They may or they may not have perennial
roots. In trees and shrubs, also in
herbs with growth from year to } r ear
from a strong tap-root, the root
is naturally perennial. But in most
perennials with only fibrous roots,
these are produced anew from time
to time or from year to year. Also,
while some such roots remain fibrous
and serve only for absorption, others
ma3 T thicken in the manner of the
ordinary biennial root and serve a
similar use, i. e. become reservoirs of
elaborated nourishment. The Dahlia
(Fig. 68) and the Peony afford good
examples of this. Sweet potato is
another instance. 1 Most such roots
have only a biennial duration : they
are produced in one growing season ; they yield their store to
form or aid the growth of the next. When perennials store up
nutritive matter underground, the deposit is more commonly
made in a subterranean portion of the stem, in tubers, corms,
bulbs, &c. (See 115-122.)
56. The distinction between annuals and biennials is at times
so difficult, and the particular in which the} T agree so manifest,
namely, that of blossoming only once, then dying, as it were
by exhaustion, that it was proposed by DeCandolle to unite
1 It is only by the readiness of this root to produce adventitious buds,
especially from its upper part, that it has been mistaken for a tuber, such
as the common potato.
FIG. 68. Fascicled and tuberous or fusiform (secondary) roots of Dahlia: a, a. buds
on base of the stem.
MORPHOLOGY OF THE ROOT. 33
the two under the common appellation of MONOCARPIC plants,
Plantte monocarpicce , taken in the sense of only once-fruiting
plants ; and to designate perennials by the corresponding term
of POLYCARPIC, Plantce poly earpiece, literally many-fruited, taken
in the sense of many-times fruiting. 1
57. But the distinction even here is no more absolute than
that between annuals and biennials. For example, it is not
quite clear whether the Cardinal Flower and related species of
Lobelia should be ranked as annuals, biennials, or perennials.
The plants may blossom and seed toward the end of the season
in which they came from seed ; or, germinated in autumn, the
small seedlings may survive the winter ; but whenever fructified
the fibrous-rooted mother plant dies throughout ; yet usually not
before it has established, and perhaps detached from the base,
small offsets to blossom the next season ; and so on. Then
Houseleeks (Sempervivum) and such-like fibrous-rooted succu-
lent plants multiply freely by offsets which are truly perennial
in the sense that they live and grow for a few or several years ;
but when at length a flowering stem is sent up producing blos-
som and seed, that plant dies as completely and in the same
manner as any biennial, only the generation of offsets surviving.
The same is true of the Century plant (Agave Americana,
wrongly denominated American Aloe) , which vegetates in the
manner of the accumulating stage of a biennial, except that
this continues for several or very many years, while the flower-
ing stage, when it arrives, is precipitated and terminated in a
single season.
58. Although the stem usually sends forth roots only when
covered by or resting on the soil, which affords congenial dark-
ness and moisture, yet these are in some cases produced in the
open air. Roots may likewise subserve other and more special
uses than the absorption of crude or the storing of elaborated
nourishment.
59. Aerial Roots is a general name for those which are pro-
duced in the open air. One class of these may serve the office
of ordinary roots, by descending to the ground and becoming
established in the soil. This occurs, on a small scale, in the
stems of Indian Corn ; the lower nodes emitting roots which
grow to the length of several inches before they reach the ground
1 These terms or some equivalents have a convenience in descriptive
botany. But those employed by DeCandolle are not happily chosen, as has
often been said. Montftocons (bearing progeny once) and Polytocoiis (bearing
many times) would be more appropriate.
3
34
MORPHOLOGY OF THE KOOT.
into which the}* penetrate. More remarkable cases abound in
those tropical regions where the sultry air, saturated with moist-
ure for a large part of the year, favors the utmost luxuriance of
vegetation. In the Palm-like Fandanus or Screw-Pine l (Fig.
69), very strong roots, emitted in the open air from the trunk,
and soon reaching
the soil, give the
appearance of a tree
partially raised out
of the ground. The
famous Banyan -tree
of India (Fig. 71) is
a still more striking
illustration ; for the
aerial roots strike
from the horizontal
branches of the tree,
often at a great
height, at first swing-
ing free in the air,
but finally reach-
ing and establishing
themselves in the
ground, where the}^
increase in diameter
and form accessory
trunks, surrounding
the original bole and supporting the wide-spread canopy of
branches and foliage. Veiy similar is the economy of the Man-
grove (Fig. 70), which forms impenetrable thickets on low and
muddy sea-shores in the tropics throughout most parts of the
world, extending even to the coast of Florida and Louisiana.
Here aerial roots spring not only from the main trunk, as in
the Pandanus, but also from the branchlets, as in the Banyan.
Even the radicle of the embryo starts into growth, protrudes,
and attains considerable length while the fruit is still attached to
the branch.
59. Aerial Rootlets for climbing are familiar in the Ivy of the
Old World (Heclera), Trumpet-Creeper (Tecoma radicans) , and
our Poison Ivy (Rhus Toxicodendron) ; by the adhesion of
1 So named, not from any resemblance to a Pine-tree, but from a like-
ness of the foliage to that of a Pine-Apple.
FIG. 69. Pandanus, or Screw- Pine; and in the background, 70, a Mangrove-tree
(Rhizophora Mangle).
MORPHOLOGY OF THE KOOT. 3b
which the stems, as they grow, ascend walls and the trunks of
trees with facilit}'. In Rhus a superabundance of these rootlets
is produced, thickly covering all sides of the stem.
60. Epiphytes or Air-Plants also have roots which are through-
out life unconnected with the ground. Epiphytes, or Epiphytic
plants, as the name denotes, are such as grow upon other plants
without taking nourishment from them. Deriving this from the
air alone, the} r are called Air-plants. This name might be
extended to the same or other kinds of plants attaching them-
selves to bare walls, rocks, and the like, and unconnected with
the soil, though such would not technically be epiphytes. Very
many Lichens, Mosses, and other plants of the lower grade, and
not a few phsenogamous plants, are in this case. The greater
part of the pliomogamous Epiphytes pertain to two monocotyle-
donous orders, the Orchis famil}' and that to which the Pinc-
Apple belongs, viz. the Bromeliaceae. Their thread-like or
cord-like simple roots cither adhere to the bark of the supporting
tree, securing the plant in its position, or some hang loose in the
air. Of these, Orchids, i. e. plants of the Orchis family, are the
most show}' and numerous, and of the greatest variety of forms,
especially of their blossoms, which are often bizarre and fantas-
tic. They belong, naturally, to climates which are both warm
and humid ; they arc highly prized in hot-house cultivation ;
and, along with the hardy and terrestrial portion of the order,
they are peculiarly interesting to the botanist on account of the
singular and exquisite adaptation of their flowers in relation
to insects which visit them. In some the blossoms curiously
FIG. 71. The Banyan-tree, or Indian Fig (Ficus Imllca),
36 MORPHOLOGY OF THE BOOT.
resemble butterflies or other insects ; as, for example, Oncidium
Papilio, Fig. 72. Epiphytic orchids are indigenous to the United
States only from Georgia to Texas, and only in humble forms,
in company with species of Tillandsia, representing Bromeliace-
ous epiphytes. The commonest of the latter tribe, and of most
northern range, is the T. usneoides, the so-called Long Moss,
which, pendent in long and tangled gray clusters or festoons from
the branches of the Live-Oak or Long-leaved Pine, gives such a
peculiar and sombre aspect to the forests of the warmer portions
of our Southern States.
61. Parasitic Plants have the peculiarity that their roots, or
what answer to roots, not only fix themselves to other plants,
but draw therefrom their nourishment, at least in part. Among
ciyptogamous plants very many Fungi are parasitic upon or
within living plants or animals. But onlj' phaenogamous para-
sites are here under consideration. These ma} r be divided into
two classes ; those with and those without green foliage.
62. Green Parasites may be either wholly or partially parasitic ;
that is, they may draw all their support from a foster plant, or
FIG. 72. Oncidium Papilio, and, 73. Comparettia rosea; two showy epiphytes of the
Orchis family; showing the mo) to the epidermis (e) of a stem of
Negnndo, a year old : B. the bark; W. the wood; and C. the cambium-layer, as found
in February. The parts referred to by small letters are: p. a portion of the pith:
mr. small portion of a medullary ray where it runs into the pith; four complete med-
ullary rays as seen on a transverse section, appear in the upper figure, running from
pith to bark : ms. medullary sheath, a circle of spiral unreliable ducts, one seen length-
wise with uncoiling extremity in the lower figure: w, w. woody tissue: dd. one of the
dotted ducts interspersed in the wood: cl. cambium-layer or zone of new growth of
wood and inner bark : l-b. liber or inner bark, the inner portion of which is here cellular,
the outer (ft) composed of slender and thick-walled bast-cells or true liber-cells:
f/e. green envelope or inner cellular bark: ce. corky envelope or outer cellular bark:
e. epidermis.
78 MORPHOLOGY OF STEMS.
their strength and toughness. They are like wood-cells except
in their greater length and flexibility, and in the thickness of
their walls, which greatly exceeds the calibre. This is the
material which gives to the bast or inner bark of Basswood, &c.,
the strength and pliability that adapts it for cordage and for
making mats : it is the material of linen, and the like textile
fibres. (For a view of the whole composition and structure of a
woody stem at the close of the first year's growth, and immedi-
ately before that of the second year begins, see Fig. 135 a .)
143. Annual Increase in Diameter. An herbaceous stem does
not essentially differ from a woody one of the same age, except
that the wood forms a less compact or thinner zone ; and the
whole perishes, at least down to the ground, at the close of the
season. But a woody stem makes provision for continuing its
growth from year to year. As the layer of wood continues to
increase in thickness throughout the season, by the multiplication
of cells on its outer surface, between it and the bark, and when
growth ceases this process of cell-multiplication is merely sus-
pended, so there is always a zone of delicate }~oung cells in-
terposed between the wood and the bark. This is called the
CAMBIUM, or, better, the CAMBIUM-LAYER. It is charged with
organizable matter, which is particularly abundant and mucila-
ginous in spring when growth recommences. This mucilaginous
matter was named Cambium by the older botanists : they sup-
posed as is still popularly thought that the baik, then so
readily separable, really separated from the wood in spring, that a
quantity of rich mucilaginous sap was poured out between them
and became organized into a tissue, the inner part becoming new
wood, the outer, new bark. But delicate slices show that there
is then no more interruption of the wood and inner bark than
at any other season. The bark, indeed, is then very readily
detached from the wood, because the cambium-layer is gorged
with sap ; but such separation is effected by the rending of a
delicate forming tissue. And if some of this apparent mucilage
be scraped off from the surface of the wood, and examined under
a good microscope, it will be seen to be a thin stratum of young
wood-cells, with the ends of medullary rays here and there in-
terspersed. The inner portion of the cambium-layer is therefore
nascent wood, and the outer is nascent bark. As the cells of
this layer multiply, the greater number lengthen vertically into
woody tissue : some are transformed into ducts ; and others,
remaining as parenchyma, continue the medullary rays or com-
mence new ones. In this wa}*, a second layer of wood is formed
the second season over the whole surface of the former layer
INTERNAL STRUCTURE. 79
between it and the bark ; and this is continuous with the woody
la}*er of the new roots below and of the leafy shoots of the sea-
son above. Each succeeding } T ear another la} T er is added to the
wood in the same manner, coincident with the growth in length
by the development of the buds. A cross-section of an exoge-
nous stem, therefore, exhibits the wood disposed in concentric
rings between the bark and the pith ; the oldest lying next the
latter, and the youngest occupj'ing the circumference. Each
layer being the product of a single year's growth, the age of an
exogenous tree ma}*, in general, be correctly ascertained by
counting the rings in a cross-section of the trunk. 1
144. Demarcation of the Annual Layers results from two or more
causes, separate or combined. In oak and chestnut wood, and
the like, the layers are strongly defined by reason of the accumu-
lation of the large dotted ducts (here of extreme size and in
great abundance) in the inner portion of each hiyer, where their
open mouths on the cross-section are conspicuous to the naked
e}*e, making a strong contrast between the inner porous and the
exterior solid part of the successive layers. In maple and beech
wood, however, tne ducts are smaller, and are dispersed through-
out the whole breadth of the layer; and in coniferous wood, viz.
that of Pine, C}*press, &c., there are no ducts at all, but only a
uniform wood}' tissue of a peculiar sort. In all these, the de-
marcation between two la} T ers is owing to the greater fineness of
the wood-cells formod at the close of the season, viz. those at
the outer border of the layer, while the next layer begins, in its
1 The annual layers are most distinct in trees of temperate climates like
ours, where there is a prolonged period of total repose, from the winter's
cold, followed by a vigorous resumption of vegetation in spring. In tropical
trees, they are rarely so well defined ; but even in these there is generally a
more or less marked annual suspension of vegetation, occurring, however,
in the dry and hotter, rather than in the cooler season. There are numerous
cases, moreover, in which the wood forms a uniform stratum, whatever be
the age of the trunk, as in the arborescent species of Cactus ; or where the
layers are few and by no means corresponding with the age of the trunk, as
in the Cycas.
In many woody climbing or twining stems, such as those of Clematis,
Aristolochia Siplio, and Menispermum Canadense, the annual layers are
rather obscurely marked, while the medullary rays are unusually broad ;
and the wood, therefore, forms a series of separable wedges disposed in a
circle around the pith. In the stem of Bignonia capreolata, the annual rings,
after the first four or five, are interrupted in four places, and here as many
broad plates of cellular tissue, belonging properly to the bark, are inter-
posed, passing at right angles to each other from the circumference towards
the centre, so that the transverse section of the wood nearly resembles a
Maltese cross. But these are exceptional cases, which scarcely require
notice in a general view.
80 MORPHOLOGY OF STEMS.
vigorous vernal growth, with much larger cells, thus marking
an abrupt transition from one la}'er to the next. Besides being
finer, the later wood-cells of the season are commonly flattened
antero-posteriorly, probably b}^ growing under greater pressure.
145. Each layer of wood, once formed, remains essentially
unchanged in position and dimensions. But, in trunks of con-
siderable age, the older layers undergo more or less change in
color, density, perviousness to moisture, &c.
146. Sap-wood (ALBURNUM). In the plantlet and in the
developing bud, the sap ascends through the whole tissue, of
whatever sort : at first through the parenchyma, for there is then
no other tissue ; and the transmission is continued through it,
especially through its central portion, or the pith, in the growing
apex of the stem throughout. But, in the older parts below, the
pith, soon drained of sap, becomes filled with air in its place,
and thenceforth it bears no part in the plant's nourishment. As
soon as wood-cells and ducts are formed, they take an active
part in the conveyance of sap, for which their tubular and ca-
pillary character is especially adapted. But, the ducts in older
parts, except when gorged with sap, contain air alone ; and in
woody trunks the sap continues to rise year after year to the
places where growth is going on, mainly through the proper
woody tissue of the wood. In this transmission, the new layers
are most active ; and these are in direct communication with the
new roots on the one hand and with the buds or shoots and leaves
of the season on the other. So, by the formation of new annual
layers outside of them, the older ones are each year removed a
step farther from the region of growth ; or rather the growing
stratum, which connects the fresh rootlets that imbibe with the
foliage that elaborates the sap, is each year removed farther from
them. The latter, therefore, after a few years, cease to convey
sap, as they have long before ceased to take part in any vital
operations. The cells of the older layers, also, usually come to
have thicker walls and smaller calibre than those of the newer.
Thus arises a distinction sometimes obscurely marked, some-
times abrupt and conspicuous into sap-wood and hcan-wood.
The former is the popular name given to the outer and newer
la}'ers of softer, more open, and bibulous wood. The early physi-
ologists named it alburnum from its white or pale color. Being
more or less sappy, or containing soluble organic matter, and
readily imbibing moisture, this part of the wood is liable to decay,
and it is therefore discarded from timber used for construction.
147. Heart-wood (or DURAMEN, so called from its greater hard-
ness or durability) is the older and mature portion of the wood.
INTERNAL STRUCTURE. 81
In all trees which have the distinction between the sap-wood and
heart- wood well marked, the latter acquires a deeper color, and
that peculiar to the species, such as the dark brown of the Black
Walnut, the blacker color of the Ebon}^ the purplish-red of Red
Cedar, and the bright yellow of the Barberry. These colors are
owing to special vegetable products, or sometimes to alterations
resulting from age. In the Red Cedar, the deep color belongs
chiefly to the medullary rays. In many of the softer woods, there
is little change in color of the heart-wood, except from incipient
decay, as in the White Pine, Poplar, Tulip-tree, &c. The
heart- wood is no longer in any sense a living part : it may perish,
as it frequently does, without affecting the life or health of the
tree.
148. The Growth and Duration of the Bark, also the differences
in structure, are much more various than of the wood. Moreover,
the bark is necessarily subject to grave alterations with advanc-
ing age, on account of its external position ; to distention from
the constantly increasing diameter of the stem within, and to
abrasion and decay from the influence of the elements without.
It is never entire, therefore, on the trunks of large trees ; but
the dead exterior parts, no longer able to enlarge with the en-
larging wood, are gradually fissured and torn, and crack off in
strips or pieces, or disappear by slow decay. So that the bark
of old trunks bears only a small proportion in thickness to the
wood, even when it makes an equal amount of annual growth.
149. The three parts of the bark (142), for the most part
readily distinguishable in the bark of young shoots, grow inde-
pendently, each by the addition of new cells to its inner face, so
long as it grows at all. The green layer commonly does not
increase after the first year ; the opaque corky layer soon excludes
it from the light ; and it gradually perishes, never to be renewed.
The corky layer usually increases for a few j'ears only, by the
formation of new tabular cells : occasionally it takes a remarkable
development, forming the substance called Cork, as in the Cork
Oak, and the thin and parchment-like layers of the White and
Paper Birches.
150. The liber, or inner bark, continues its growth through-
out the life of the exogenous tree, by an annual addition from
the cambium-layer applied to its inner surface. Sometimes this
growth is plainly distinguishable into layers, corresponding with
or more numerous than the annual layers of the wood : often,
there is scarcely any trace of such layers to be discerned. In
composition and appearance, the liber varies greatly in different
plants, especially in trees and shrubs. That of Basswood or
82 MORPHOLOGY OF STEMS.
Linden, and of other plants with a similar fibrous bark, may be
taken as best representing the liber. Here it consists of alter-
nate strata of fibrous bast, and of the peculiar liber-cells called
sieve-cells, in which nourishing matter is especially contained
and elaborated. While the latter, or their equivalents, occur
and play an important part in all inner bark, the bast-cells are
altogether wanting in the bark of some plants, and are not pro-
duced after the first 3'ear in many others. The latter is the case
in Negundo, where abundant bast-cells, like those of Basswood,
compose the exterior portion of the first year's liber, but none
whatever are formed in the subsequent layers. In Beeches and
Birches, also, a few bast-cells are produced the first }'ear, but
none afterwards. In Maples, a few are formed in succeeding
years. In the Pear, bast-cells are annually formed, but in very
small quantity, compared with the parenchymatous part of the
liber. In Pines, at least in White Pines, the bark is nearly as
homogeneous as the wood, the whole liber, except what answers
to the medullary rays, consisting of one kind of cells, resembling
those of bast or of wood in form, but agreeing with the proper
liber-cells in their structure and markings.
151. The bark on old stems is constantly decaying or falling
away from the surface, without airy injui\y to the tree ; just as
the heart-wood within may equally decay without harm, except
by mechanically impairing the strength of the trunk. There are
great differences as to the time and manner in which the older
bark of different shrubs and trees is thrown off. Some have
their trunks invested with the liber of man}^ years' growth,
although only the innermost layers are alive ; in others, it scales
off much earlier. On the stems of the common Hone} T suckle, of
the Nine-Bark (Spiraea opulifolia) , and of Grape-vines (except
Vitis vulpina) , the liber lives only one season, and is detached
the following year, hanging loose in papery layers in the former
species, and in fibrous shreds in the latter.
152. While the newer laj'ers of the wood abound in crude sap.
which the}' convey to the leaves, those of the inner bark abound
in elaborated sap, which the}' receive from the leaves and convey
to the cambium-la3*er or zone of growth. The proper juices and
peculiar products of plants are accordingly found in the foliage
and the bark, especially in the latter. In the bark, therefore
(either of the stem or of the root) , medicinal and other principles
are usually to be sought, rather than in the wood. Nevertheless,
as the wood is kept in connection with the bark by the medullary
rays, many products which probably originate in the former are
deposited in the wood.
INTERNAL STRUCTURE. 83
153. The Living Parts of a tree or shrub, of the exogenous
kind, are obviously only these: 1st, The summit of the stem
and branches, with the buds which continue them upwards and
annually develop the foliage. 2d, The fresh roots and rootlets
annually developed at the opposite extremity. 3d. The newest
strata of wood and bark, and especially the interposed cambium-
layer, which, annually renewed, maintain a living communication
between the rootlets on the one hand and the buds and foliage
on the other, however distant they at length may be. These are
all that is concerned in the life and growth of the tree ; and these
are annually renewed. The branches of each year's growth are,
therefore, kept in fresh communication, by means of the newer
layers of wood, with the fresh rootlets, which are alone active in
absorbing the crude food of the plant from the soil. The fluid
they absorb is thus conveyed directly to the branches of the sea-
son, which develop leaves to digest it. And the sap they receive,
having been elaborated and converted into organic nourishing
matter, is partly expended in the upward growth of new branches,
and partly in the formation of a new layer of wood, reaching
from the highest leaves to the remotest rootlets.
154. Longevity of trees. As the exogenous tree, therefore,
annually renews its buds and leaves, its wood, bark, and roots,
every thing, indeed, that is concerned in its life and growth,
there seems to be no necessary cause, inherent in the tree itself,
why it may not live indefinitely. Some trees are known to have
lived for one and two thousand }'ears, and some are possibly
older. 1 Equally long may survive such endogenous trees as the
Dragon tree (Dracaena) , which have provision for indefinite in-
crease in diameter (138), and for the production of branches.
The famous Dragon tree of Orotava, in Teneriffe, now destroyed
by hurricanes and other accidents, had probably reached the age
of more than two thousand years.
155. On the other hand, increase in height, spread of branches
and length of root, and extension of the surface over which the
annual layer is spread, are attended with inevitable disadvantage,
which must in time terminate the existence of the tree in a way
quite analogous to the death of aged individual animals, which
is not directly from old age, but from casualties or attacks to
1 The subject of the longevity of trees has been discussed by DeCandolle,
in the "Bibliotheque Universelle" of Geneva, for May, 1831, and in the second
volume of his " Physiologic Ve'ge'tale ; " more recently, by Alphonse DeCan-
dolle in the " Bibliotheque Universelle ; " and in this country by myself hi the
'* North American Review," for July, 1844. For an account of the huge ^Icd-
woods (Sequoias) of California, see Whitney's Yosemite Book.
84 MORPHOLOGY OF STEMS.
which the aged are either increasingly incident or less able to
resist. A tree like the Banyan (59, Fig. 71), which by aerial
roots continues to form new trunks for the support and sustenance
of the spreading branches, and thus ever advances into new soil,
has a truly indefinite existence ; but, then, it becomes a forest,
or is to be likened to a colony propagated and indefinitely in-
creased by suckers, offsets, or other subterranean shoots. So
the question of the secular continuation of the individual plant
becomes merged in that of continuation of the race, at least of
a bud-propagated race, the answer to which is wholly in the
domain of conjecture. 1 However this may be, it is evident that
a vegetable of the higher grade is not justly to be compared with
an animal of higher grade ; that individual^* is incomplete ly
realized in the vegetable kingdom ; 2 that rather
156. The Plant is a Composite Being, or Community, lasting, in
the case of a tree, through an indefinite and often immense num-
ber of generations. These are successively produced, enjoy a
term of existence, and perish in their turn. Life passes onward
continually from the older to the newer parts, and death follows,
with equal step, at a narrow interval. No portion of the tree is
now living that was alive a few years ago ; the leaves die annu-
ally and are cast off, while the internodes or joints of stem that
bore them, as to their wood at least, buried deep in the trunk
under the wood of succeeding generations, are converted into
lifeless heart-wood, or perchance decayed, and the bark that
belonged to them is thrown off from the surface. It is the aggre-
gate, the blended mass alone, that long survives. Plants of
single cells, and of a definite form, alone exhibit complete indi-
viduality ; and their existence is extremely brief. The more
complex vegetable of a higher grade is not to be compared with
the animal of the highest organization, where the offspring always
separates from the parent, and the individual is simple and indi-
visible. But it is truly similar to the branching or arborescent
coral, or to other compound animals of the lowest grade, where
successive generations, though capable of living independently
and sometimes separating spontaneously, yet are usually devel-
oped in connection, blended in a general body, and nourished
more or less in common. Thus, the coral structure is built up
by the combined labors of a vast number of individuals, by
the successive labprs of many generations. The surface or the
recent shoots only are alive ; beneath are only the dead remains
1 See Darwiniana, xii. 338-355.
2 As, perhaps, was first explicitly stated by Engelmann, in his inaugural
essay, De Antholysi Prodromus, Introduction, 4.
MORPHOLOGY OF LEAVES. 85
of ancestral generations. As in a genealogical tree, only the
later ramifications are among the living. The tree differs from
the coral structure in that, as it ordinarily imbibes its nourish-
ment mainly from the soil through its roots, it makes a downward
growth also, and. by constant renewal of fresh tissues, maintains
the communication between the two growing extremities, the
buds and the rootlets. Otherwise, the analogy of the two, as to
individuality, is well-nigh complete.
SECTION IV. OF LEAVES.
1. THEIR NATURE AXD OFFICE.
157. Leaf (Lat. Folium, in Greek form Phyllum), as a botani-
cal term, has on the one hand a comprehensive, on the other a
restricted sense. In its commonest sense, as used in descriptive
botany, it denotes the green blade only. Yet it is perfectly
understood that the footstalk is a part of the leaf, and therefore
that the phrase " leaves cordate," or the like, is a short way of
saying that the blade of the leaf is cordate or heart-shaped.
Moreover, two appendages, one on each side of the base of the
footstalk, when there is any, are of so common occurrence that
they are ranked as a proper part of the organ. So that, to the
botanist, a typical leaf consists of three parts: 1, BLADE or
LAMINA ; 2, FOOT-STALK or LEAF-STALK, technically PETIOLE ;
3, A pair of STIPULES. (Fig. 142.)
158. The blade, being the most important part of an ordinary
leaf, may naturally be spoken of as the whole. Petiole and
stipules are indeed subsidiary when present, and are not rarely
wanting. Yet sometimes they usurp the whole function of foli-
age, and sometimes there is no such distinction of parts.
159. Physiologically, leaves are green expansions borne by the
stem, outspread in the air and light, in which assimilation (3)
and the processes connected with it are earned on. Vegetable
assimilation, the most essential function of plants, being the
conversion of inorganic into organic matter, takes place in
all ordinary vegetation only in green parts, and in these when
exposed to the light of the sun. And foliage is an adaptation
for largel}* increasing the green surface. But stems, when green,
take part in this office in proportion to the amount of surface,
sometimes monopolize it, and in various cases increase their
means of doing so b}' assuming leaf-like forms. (126-129.)
Leaves, especially in such cases, may lose this function, appear
only as useless vestiges, or may be subservient to various wholly
86 MORPHOLOGY OF LEAVES.
different uses. Form and function, therefore, are not sure indi-
cations of the true nature of organs.
160. Morphologically, and in the most comprehensive sense,
leaves are special lateral outgrowths from the stem, definitely
and symmetrically arranged upon it ; in ordinary vegetation and
in the most general form constituting the assimilating apparatus
(or foliage) , but also occurring in other forms and subserving
various uses. Sometimes these uses are combined with or sub-
sidiary to the general function of foliage ; sometimes the leaf is
adapted to special uses only. So the botanist recognizing the
essential identity of organs, whatever their form, which appear
in the position and conform to the arrangement of leaves
discerns the leaf in the cotyledons of a bean or acorn, the scale of
a lily-bulb or the coat of an onion, the scale of a winter bud, and
the petal of a blossom. Therefore, while expanded green leaves
(which may be tautologically termed foliage-leaves) are taken as
the proper t}*pe, the common name of leaves, in the lack of any
available generic word, is in morphological language extended to
these special forms, whenever it becomes needful to express their
phylline or foliar nature.
161. In the morphological view, all the plant's organs except-
ing roots (and excepting mere superficial productions, such as
hairs, prickles, &c.), belong either to stem or to leaves, are
either cauline or phylline in nature. To the latter belong all the
primary outgrowths from nodes, all lateral productions which
are not axillary. 1 Whatever is produced in the axil of a leaf is
cauline, and when developed is a branch.
162. The Duration of Leaves is transient, compared with that
of the stem. They may be fugacious, when they fall off soon
after their appearance ; deciduous, when they last only for a
single season ; and persistent, when the}' remain through the cold
season, or other interval during which vegetation is interrupted,
and until after the appearance of new leaves, so that the stem is
never leafless, as in Erergreens. In many evergreens, the leaves
have onh' an annual duration ; the old leaves falling soon after
those of the ensuing season are expanded, or, if they remain
longer, ceasing to bear any active part in the economy of the
vegetable, and soon losing their vitality altogether. In Pines
and Firs, however, although there is an annual fall of leaves
either in autumn or spring, }*et these were the produce of some
1 There are cases in which this rule is of difficult application, or is seem-
ingly violated, sometimes by the suppression of the subtending leaf, as in the
inflorescence of Cruciferse, rarely in other ways, to be explained in the
proper places.
THEIR STRUCTURE AND FORMS. 87
season earlier than the last ; and the branches are continually
clothed with the foliage of from two to five, or even ten or more
successive years. On the other hand, it is seldom that all the
leaves of an herb endure through the whole growing season, the
earlier foliage near the base of the stem perishing while fresh
leaves are still appearing above. In our deciduous trees and
shrubs, however, the leaves of the season are mostly developed
within a short period, and they all perish in autumn nearly
simultaneously.
1G3. Leaves soon complete their growth, and have no power
of further increase. Being organs for transpiration, a very large
part of the water imbibed by the roots is given out by the foliage,
leaving dissolved earthy matters behind. Assimilation can take
place only in fresh and vitally active tissue. It is incident to all
this that leaves should be of only transient duration, at least in
their active condition.
164. Defoliation. The leaves of most Dicotyledons and some
Monocotyledons separate from the stem and fall by means of an
articulation at the junction with the stem, which begins to form
earl}' in the season and is completed at the close. There is a kind
of disintegration of a transverse layer of cells, which cuts off the
petiole by a regular line, and leaves a clean scar, such as is seen
in Fig. 81, 85, 91. Some leaves, notably those of Palms,
Yucca, and other endogens, die and wither on the stem, or wear
away without falling.
165. In temperate climates, defoliation mostly takes place at
the approach of winter. In warmer climates having only winter
rain, this occurs in the hot and dry season.
166. Normal Direction or Position. The leaf-blade is expanded
horizontally, that is, has an upper and an under surface. When
erect, the upper surface faces the axis which bears it. To this,
there are many seeming but no real exceptions ; that is, none
which are not explicable as deviations or changes from the normal
condition. (213-217.)
2. THEIR STRUCTURE AND FORMS AS ORGANS OF ASSIMILATION
OR VEGETATION, i. e. AS FOLIAGE.
167. The Internal Structure or Anatomy of the leaf needs here
to be examined so far as respects its obvious parts and their
general composition. The leaf, like the stem, is composed of
two elements (131), the cellular and the woody. The cellular
portion is the green pulp or parenchyma, and in this the work
of assimilation is carried on. The woody is the fibrous frame-
88 MORPHOLOGY OF LEAVES.
work, the separate parts or ramifications of which form what are
variously called the ribs, a sufficiently proper term, nerves or
veins. The latter names ma}- suggest false analogies ; but they
are of the commonest use in descriptive botan}*. That of veins,
and of its diminutive, veinlets, for the smaller ramifications, is
not amiss ; for the fibrous framework not only gives firmness
and support to the softer cellular apparatus, i. e. forms ribs,
but serves in the leaf, as it does in the stem, for the more rapid
conveyance and distribution of the sap. The subdivisions con-
tinue bej'ond the limits of unassisted vision, until the fibro-vas-
cular bundles are reduced to attenuated fibres ramified through
the parenchyma. In leafstalks, the woody bundles are parallel,
not ramified, and arranged in various ways ; in Exogens usually
so as to form in cross-section an arc or an incomplete or com-
plete ring. In leaves serving as foliage or organs of assimilation,
the blade is the important part, and this only is here regarded.
168. The characteristic contents of these cells of parenchyma
are grains of chlorophyll (142 3 ), literally leaf-green, to which
the green color of foliage is wholly owing, and which may be
regarded as the most important of all vegetable products ;
because it is in them (or in this green matter, whatever its form,
189) that all ordinary assimilation takes place. As it acts only
under the influence of light, the expanded leaf-blade may be
viewed as an arrangement for exposing the largest practicable
amount of this green matter the essential element of vegeta-
tion to the light and air.
169. The Parenchyma-cells, constituting the green pulp, are
themselves arranged in accordance
with this adaptation. The upper stra-
tum is mostly of oblong cells, compactly
arranged in one or more layers, their
longer diameter perpendicular to the
surface. The stratum next the lower
surface of the leaf consists of loosely
arranged cells, with longer diameter
usually parallel to the plane of the leaf,
133 often irregular in form, and so disposed
as to leave intervening sinuous air-spaces freely permeating all
FIG. 136. A magnified section through the thickness of a leaf of lllidum Flori-
danum, showing (ho irregular spaces or passages between the cells, which are small in
the upper layer of the green pulp, the cells of which (placed vertically) are well com-
pacted, so as to leave only minute vacuities at their rounded ends; but the spaces are
large and copious in the rest of the leaf, where the cells are very loosely arranged :
also the epidermis or skin of the upper (a) and of the lower surface of the leaf (b\
composed of irfectly combined and thick-walled empty cells.
THEIR STRUCTURE AND FORMS.
89
that part of the leaf. (Fig. 136, 137.) Hence in good part the
deeper green hue of the upper, and the paler of the lower face
of leaves.
170. Epidermis. The whole surface of leaves, as of young
stems, is invested with a translucent membrane, composed of
one or sometimes two or three layers of empty and rather-
thick-walled cells. This is the skin or epidermis, which is so
readily separable from the succulent tissue of such leaves as
those of Stonecrop and other species of Sedum. It is of a single
layer in the Illicium (Fig. 136) and Lily (Fig. 137) ; of as
man)' as three in the firm leaf of the Oleander ; is generallj*
hard and thick in such coriaceous leaves as those of Pittosporum
and Laurustinus, which thereby the better endure the dry air of
rooms in winter.
171. Stomata or Breathing-pores. 1 The epidermis forms a
continuous protective investment of the leaf except where certain
organized openings occur, the stomata. They are formed by a
transformation of some of the cells of the epidermis ; and consist
usually of a pair of cells (called guardian-cells) , with an opening
between them, which communicates with an air-chamber within,
and thence with the irregular intercellular spaces which permeate
the interior of the leaf. Through the stomata, when open> free
interchange may take place between the external air and that
1 The technical name has been anglicized stomates, singular stomate, which
has no advantage over the proper Greek, sing, sfoma, pi. stomata.
FIG. 137. A magnified section through the thickness of a minute piece of the leaf
of the White Lily of the gardens, showing also a portion of the under side with some
breathing-pores, stoinata.
90
MORPHOLOGY OF LEAVES.
within the leaf, and thus transpiration be much facilitated.
When closed, this interchange will be interrupted or impeded.
The mechanism of stomata is somewhat
recondite, and will be illustrated in the
anatomical and physiological volume of
this series.
172. It is only when leaves assume
a vertical or edgewise position that the
stomata are in equal numbers on both
faces of a leaf. Ordinarily, the y occupy
or most abound on the lower face, which
is turned away from the sun ; but in certain coniferous trees the
reverse of this is true. In the Water Lilies (Nymphsea, Nuphar),
and other leaves which float
upon the water, the stomata all
belong to the upper surface.
Leaves which live under water,
where there can be no evapora-
tion, are destitute, not only of
stomata, but usually of a distinct epidermis also. The number
of the stomata varies from 800 to about 170,000 on the square
inch of surface in different leaves. In the Apple, there are said
to be about 24,000 to the square inch (which is under the average
number, as given in a table of 36 species by Lindley) ; so that
each leaf of that tree would present about 100,000 of these
orifices. The leaf of Dragon Arum is said to have 8,000
stomata to a square inch of the upper surface, and twice that
number in the same space of the lower. That of the Coltsfoot
has 12,000 stomata to a square inch of the lower epidermis,
and only 1,200 in the upper. That of the White Lily has
from 20,000 to 60,000 to the square inch on the lower sur-
face, and perhaps 3,000 on the upper; and they are so re-
markably large that they may be discerned Ly a simple lens of
an inch focus.
172. Venation, the veining of leaves, &c., relates to the mode
in which the woody tissue, in the form of ribs, veins, &c., is
distributed in the cellular. There are two principal modes, the
parallel-veined and the reticulated or netted-veined. The former
is especially characteristic of plants with endogenous stem and
monocotyledonous embryo, and also of gymnospermous trees,
FIG. 138. A highly magnified piece of the epidermis of tlie Garden Balsam, with
three stomata (after Brongniart).
FIG. 139. Magnified view of the 10,000th part of a square inch of the epidermis of
the lower surface of the leaf of the White Lily, with its stomata. 140. A single stoma,
more magnified. 1 11. Another stoma, widely open.
THEIR STRUCTURE AND FORMS.
9.1
which have exogenous stems and at least dicotyledonous
embryos. The latter prevails in ordinary plants with exogenous
stem and dicotyledonous embryo.
173. Parallel- veined or Nerved leaves (of which Fig. 143 is
an illustration) have a framework of simple ribs (called by the
earlier botanists nerves, a name still used in descriptions), which
run from the base to tip, or sometimes from a central strong rib
to margin of the leaf, in a generally parallel and undivided way,
and sending off or connected by minute veinlets only. Grasses,
Lily of the Valley, and the like, illustrate the commoner mode
in which the threads of wood run from base to apex. The
Banana and Canna are familiar illustrations of a mode not un-
common in tropical or subtropical endogens, in which the threads
or "nerves" run from a central rib (midrib} to the margin.
Parallel- veined leaves are generally entire, or at least their
margins not toothed or indented. The principal exception to
this occurs when the ribs or the stronger ones are few in number
and radiately divergent, as in the flabelliform leaves of Fan-
palms, a peculiar modification of the parallel- veined type.
Between leaves with nerves wholly of basal origin, and those
with nerves all springing from a midrib, there are various grada-
tions, and also in respect to curving. But parallel- veined or
nerved leaves may be classified into
FIG. 142. A leaf of the Quince, of the netted-veined or reticulated sort, with
blade (6), petiole or leaf-stalk (p), and stipules (st).
FIG. 143. Parallel-veined leaf of the Lily of the Valley, Convallaria majalis.
92 . MORPHOLOGY OF LEAVES.
Basal-nerved, that is, with the nerves all springing from the
base of the leaf, and
Costal-nerved, springing from a midrib or costa. Either ma}' be
Rectinerved, the nerves running straight from origin to apex
or margin of the leaf, as the case may be ;
Currinerved, when curving in their course, as in the leaves
of Funkia and in Canna ;
Flabell inerved, where straight nerves and ribs radiate from the
apex of the petiole, as in Fan-palms and the Gingko tree.
174. In typical parallel- veined leaves, all reticulation is con-
fined to minute and straight cross-veinlets : in many, these are
coarser, branching, and reticulated ; in some, as in Smilax and
Dioscorea, only the primary ribs or strongest nerves are on the
parallel- veined plan ; the space between being filled with reticu-
lations of various strength ; thus passing by gradations into
175. Reticulated or Netted- veined leaves. In most of these,
from one to several primary portions of the framework are
particularly robust, and give origin to much more slender ram-
ifications, these to other still smaller ones, and so on. The
strong primary portions are RIBS (cosfce) ; the leading ramifi-
cations, VEINS (vence) ; the smaller and the ultimate subdivisions,
VEIXLETS (yemdce). All or some of the veins and veinlets are
said to anastomose, i. e. variously to connect with those from
other trunks or ribs, apparently in the manner of the veins and
arteries of animals, forming meshes. But, as there is no opening
of calibre of one into another, the word is etymologically rather
misleading. More properly, it is said that the veins or veinlets
form reticulations or net-work. A primary division of reticulated
leaves, and indeed of nerved leaves also, into two classes, is
founded upon the number of primary ribs.
176. There may be only a single primary rib ; this traversing
the blade from base to tip through its centre or axis (as in Fig.
142, 152-156) is called the MIDRIB. There ma}* be others, gen-
erall}* few (one, two, three, or rarely four) , rising from the apex
of the petiole on each side of the midrib, running somewhat par-
allel w r ith it or more or less diverging from it : these are lateral
ribs. Among parallel- veined leaves, the Banana, Canna, &c.,
have a single rib, from which the veins (in the older nomencla-
ture here called nerves) all proceed. Most Lilies and the like
have several approximately parallel ribs, but the midrib pre-
dominant : in other cases, the midrib is no stronger than the
others. In Fan-palms, the ribs are radiately divergent, giving
a fan-shaped or rounded outline to the blade. In reticulated
leaves, in which the veins all spring from the ribs, the two
THEIR STRUCTURE AND FORMS.
93
classes into which they divide are the pinnately veined and the
palmately veined.
177. Pinnately or Feather-reined (or Penninerved) leaves
are accordingly those of which the veins and their subdivisions
are side branches of a single central rib (midrib), which traverses
the blade from base to apex ; the veins thus being disposed in
the manner of the plume on the shaft of a feather. (Fig. 142,
152, &c.) Sometimes these continue straight and undiminished
from midrib to margins (straight- veined, as in Beech and Chest-
nut, Fig. 152), sending off only small lateral veinlets ; some-
times they ramify in their course into secondaiy or tertiary veins,
and these into veinlets. Pinnate venation in reticulated leaves
naturally belongs to leaves which are decidedly longer than wide.
178. Some of the primary veins, commonly among the lower,
may be stronger than the rest, and thus take on the character of
ribs, or by gradations pass into such. The leaf of the common
annual Sunflower (Fig. 155) becomes in this way triple-ribbed or
tripli-nerved. The appearance of a second pair of such strength
ened veins makes the venation quintupli-ribbed or quintupli-nerved.
146
15.1 154
155
Through the approximation of such strong veins to the base of
the blade, this venation may pass into the
179. Palmately, Digitately, or Radiately Veined (or Palmi-
nerved) class, of which leaves of common Maples and the Vine
are familiar examples. (Also Fig. 158-160, &c.) In these
FIG. 144-157. Various forms of simple leaves, explained in the text and in the
Glossary.
94
MORPHOLOGY OF LEAVES.
there are three, five, seven, or sometimes more ribs of equal
strength, the central being the midrib, and each with its system
of veins which ramify and form meshes in the interspaces. Here
the whole wood}- portion of the leaf divides equally into a num-
ber of parts upon leaving the petiole or entering the blade. The
ribs there commonly diverge more or less in a palmate or digitate
manner (i. e. like outspread fingers of the hand, or the claws of a
bird, or like radii of more or less of a circle) : so, in the corre-
lation of outline and venation, this class of veining goes with
158
163
164 165
roundish circumscription. This is not so true, however, in a
special case, viz., where the ribs, however divergent below, curve
forward and all run to the apex of the blade, thus imitating the
parallel-veined system, as in Rhexia and generally in the family
of which that genus is the single northern representative. 1
180. Forms as to Outline, &c. DeCandolle conceived the shape
of leaves (both the general circumscription and the special con-
figuration) to depend on the distribution of the ribs and A r eins,
and quantity of the parenchyma in which these were outspread,
a too mechanical view, and not conformable to the history of
development. This proves that the framework is adapted to the
parenchyma, which grows and shapes the organ in its own way,
rather than the parenchyma to it. It were better to say that the
1 In Linnasan terminology, palmate and digitate referred to particular out-
line only, and were separately used to denote extent of division, palmate, not
divided down to the petiole, digitate, when divided, like the claws of a bird,
quite down the base. DeCandolle generalized the use of the former term,
and ever since the two have been used interchangeably.
FIG. 158-166. Various forms of simple, chiefly palmately veined leaves.
THEIR STRUCTURE AND FORMS.
95
two elements of the structure are correlated. Descriptive terms
applied to leaves are equally applicable to all expanded organs
or parts, and indeed to all outlines. Some leading forms are
here enumerated ; and all are defined in the Glossary.
181. As to general Circumscription, proceeding from narrower
to broader shapes, and then to those with either narrowed or
notched base, leaf-blades are
167
Linear, when narrow, several times longer than wide, and of
about the same breadth throughout. (Fig. 167.)
Lanceolate, or Lance-shaped^ when several times longer than
wide, and tapering upwards (Fig. 153, 168), or tapering both
upward and downward.
Oblong, when nearly twice or thrice as long as broad. (Fig. 169.)
Elliptical, oblong with a flowing outline, the two*ends alike
in width. (Fig. 170.)
Oval, the same as broadly elliptical, or elliptical with the
breadth considerably more than half the length.
Ovate, when the outline is like a section of a hen's-egg length-
wise, the broader end being downward. (Fig. 171, 155.)
Orbicular, or Rotund, circu-
lar in outline, or nearly sp.
(Fig. 160.)
Obovate, inversely ovate, or
ovate with the narrower end
toward the base, the broader
upward. (Fig. 175, 145.)
Cuneate, or Cuneiform, that is, Wedge-shaped, broad above and
tapering by straight lines to an acute base. (Fig. 176, 148.)
Spatulate, rounded above, long and narrow below, like a
spatula. (Fig. 174, 147.)
Oblanceolate, inverted lance-shaped, i. e. such a lanceolate
leaf as that of Fig. 168, but with the more tapering end at base,
as in Fig. 173. To those who restrict the term lanceolate to the
sense of a narrow leaf tapering equally in both directions, the
17.=.
176
FIG. 167-176. Outlines of various simple leaves.
96
MORPHOLOGY OF LEAVES.
term oblanceolate is superfluous. The following terms desig-
nate leaves with a notched instead of narrowed base.
Cordate, or Heart-shaped, when a leaf of an ovate form, or
something like it, has the outline of its rounded base turned in
(forming a notch or sinus) where the stalk is attached, as in
Fig. 172, 151. Also Fig. 149, Pontederia, a leaf of the parallel-
veined class.
Reniform, or Kidney-shaped, like the last, only rounder and
i broader than long. (Fig. 158.)
Auricnlate, or Eared, having a pair of small and blunt pro-
jections, or ears, at the base, as in Magnolia Fraseri, Fig. 178.
Sagittate, or Arrow-shaped, where such ears are acute and turned
downwards, while the
main body of the blade
tapers upwards to a
point, as in the com-
mon species of Sagit-
taria or Arrow-head,
and in the Arrow-
leaved Polygonum.
(Fig. 165, 177.)
Hastate, or Halberd-
shaped, when such
lobes at the base point outwards, giving the leaf the shape of the
halberd of the olden time, as in Polygonum arifolium (Fig. 170)
and Sorrel, Fig. 163.
182. Peltate or Shield-shaped leaves are those in which a blade
of rounded or sometimes of other shape is attached to the petiole
by some part of the lower surface, instead of the basal margin :
those of Water-shield or Brasenia, of Nelumbium, and of Hydro-
cotyle umbellata are marked examples. The anomaly is mor-
phologically explained by a comparison with deepty cordate or
reniform leaves having a narrow sinus, such as those of N}'m-
phaea or Water Lily, and by supposing a union of the approxi-
mated edges of the sinus. Fig. 159 and 160, from two species
of Hydrocotyle, one with open and the other with closed sinus
obliterated by the union, illustrate this.
183. As to Extremity, whether base or apex, there are several
descriptive terms, expressive of the principal modifications ;
such as
Acuminate, tapering, either gradually or abruptly, into a
narrow more or less prolonged termination. (Fig. 180.)
FIG. 177-179. Sagittate, auriculate, and hastate leaves.
THEIR STRUCTURE AND FORMS.
97
Acute, ending in an acute angle, without special tapering, as
in Fig. 181.
Obtuse, ending with a blunt or roundish extremity, Fig. 182.
Truncate, with termination as if cut off by a straight transverse
line, as in Fig. 183.
Retuse, with an obtuse extremity slightly depressed or re-enter-
ing, as in Fig. 184.
Etnarglnate, with a more decided terminal notch, Fig. 185.
Obcordate, inversely heart-shaped, i. e. like cordate, but the
broader end and its strong notch at apex instead of base, Fig.
186. This and the following terms are applicable to apex only.
Mucronate, abruptly tipped with a small and short point, like a
projection of the midrib, as in Fig. 188.
Cuspidate, tipped with a sharp and rigid point, as in Fig. 187.
184. As to Margin or special Outline, the terminology proceeds
upon the convenient supposition of a blade with quite entire margin,
but subject to incisions, which give rise to notches or clefts, if we
regard the sinuses ; or to teeth, lobes, segments, &c., if we regard
the salient portions between the sinuses. The ribs, or the stronger
veins, &c., commonly terminate in the teeth or lobes ; but in Cicuta
maculata, and in a few other cases, they run to the notches.
185. Dentation relates to mere marginal incision, not extend-
ing deeply into the blade. The blade is said to be
Entire, 1 when
the margin is com-
pletely filled out
to an even line, as
in Fig. 173-179. j
Serrate, when j
with small and J
sharp teeth direct-)
ed forward, like \
the teeth of a saw, ^
as in Fig. 189. \N
Serrulate is the iao 190 191 192 193 194
diminutive of serrate, and is equivalent to minutety serrate.
1 Inteyerrimus-u-um, or quite entire, is the term in Latin terminology.
yer means undivided or not lobed.
FIG. 180-188. Terminations of leaves.
FIG. 189-194. Dentation of pinnately veined leaves.
7
Inte
98 MORPHOLOGY OF LEAVES.
Dentate, or Toothed, a general term for toothing, specially
applied to the case of salient teeth which are not directed for-
ward or towards the apex of the blade, Fig. 190.
Crenate, or Scalloped, the same as dentate or serrate, but with
teeth much rounded, Fig. 191.
Repand, or Undulate, when the margin is a wav} T line, bending
slightly inward and outward, Fig. 192.
Sinuate, when this wavy line is stronger or distinctly sinuous,
as in Fig. 193.
Incised, when cut by sharp and irregular incisions more or less
deeply, Fig. 194. This is intermediate between dentation and
186. Lobation or Segmentation. When the blade is more
deeply penetrated by incisions from the margin, that is, when
the spaces between the ribs or principal veins are not filled to
near the general outline, it is said to be lobed, cleft, parted, or
divided, according to the degree of separation ; and the portions
are called lobes, segments, divisions, &c. The most general name
for such parts of any simple blade is that of lobes. More par-
ticularly a leaf-blade, or other body, is said to be
Lobed, when the division extends not more than half way down,
and either the sinuses or the lobes are rounded ;
Cleft, when the division is half way down or more, and the
lobes or sinuses narrow or acute ;
Parted, when the divisions reach almost, but not quite, to the
base or the midrib ;
Divided, when the}' sever the blade into distinct parts, which
makes the leaf compound. (193.)
187. LOBE is the common name of one of the parts of a simple
blade, especially when there is only one order of incision. But
when there are more, as when a leaf is divided or parted and
these primary lobes again lobed or cleft, the lobes of first order
are commonly called SEGMENTS (sometimes divisions or partitions),
and the parts of these, Lobes. Or the lobes may be designated
as primary, secondary, tertiary, &c. Ultimate portions or small
lobes may be called Lobules or Lobelets. Also the portions of a
quite divided blade take the name of Leaflets. By proper selec-
tion of terms, the degree of division or lobing may thus be
expressed in a single word.
188. As to Number of parts, this may be tersely expressed by
combination with the adjective term applicable to the degree ; as,
Two-lobed, Three-lobed, Five-lobed, Many-lobed, &c. ; or Two-Five-
cleft, Many-cleft, &c., in Latin form Bifid, Trijid, Multifid, &c. ;
Two-jive-parted ', &c., according to the number of divisions
which extend almost to the base or axis ; Two-Five-divided (in
THEIR STRUCTURE AND FORMS.
99
Latin form Bisected, Trisected, &c.), when there are two or three
or more complete divisions of the blade.
189. As to Arrangement of parts, this may be simply and best
expressed by taking into account the nature of the venation or
the distribution of the ribs, &c., which controls or is co-ordinated
with the disposition of the lobes. Pinnately veined leaves,
when lobed, must needs have the incisions directed to the mid-
rib ; palmatery-veined or radiated, to the apex of the petiole.
The lobes or divisions of the first will be pinnately, of the second
lie
palmately disposed. Accordingly, the three leaves of as many
species of Oak, Fig. 195, 196, and 197, represent respectively a
pinnately lobed, pinnately cleft, and pinnately parted leaf, while the
accompanying leaf of Celandine, Fig. 198, is pinnately divided.
The first three, however, when the degree of incision is not par-
ticularly in question, usually pass under the common term of
pinnatifid, Fig. 195 moderately, Fig. 197 deepty. The number
of lobes, when definitely marked, may come into the descriptive
phrase, as pinnately 1 -lobed, pinnately 1 -cleft, parted, or divided,
as the case ma}- be.
190. Similarly, Figures 199 to 202 represent, respectively, a
palmately three-lobed, three-cleft, three-parted, and three-divided, or,
in Latin form, trilobate, trifid, tripartite, and trisect or trisected
leaf. Fig. 166 is & palmately ^-parted leaf; Fig. 164, palmately
FIG. 195-198. Pinnately lobed, cleft, parted, and divided leaves.
FIG. 199-1*02. Palmately 3-lobed, cleft, parted, and divided leaves.
100 MORPHOLOGY OF LEAVES.
multifid, &c. Fig. 162, a leaf of Dragon Arum, is palmntelif
^-parted. But, as the lateral sinuses are not so deep as the
others, the leaf is said to be pedately parted, or pedate, in the
earl}' terminology.
191. Moreover, as the lobes or divisions of a leaf may be
again similar!}' lobed or parted, &c., this composition ma}' be
indicated by the prefix twice, thrice, &c., as twice pinnatijid or
bipinnatifid, thrice pinnately parted, thrice palmately parted, and
the like. Thus, a word or two, or a short phrase, may describe
even a complex leaf, so as to convey a perfectly clear and defi-
nite idea of its conformation.
192. A distinction should now be drawn between simple and
compound leaves. The distinction cannot be both natural and
absolute ; for the one may pass variously into the other. Simple
leaves, which have been thus far considered, have a single lamina
or blade, which may, however, at one extreme be entire, at the
other many-parted, and even seA^eral times divided.
193. Compound Leaves are those which have from two to
many distinct blades, on a common leafstalk. These blades,
called LEAFLETS, may be sessile on the common leafstalk, or they
may have leafstalks of their own. As the leaf very commonly
separates in age by an articulation of its petiole with the stem,
so leaflets are commonly more or less articulated with the com-
mon petiole. When the leaf, with its petiole, falls from the
stem, the leaflets may as completely separate from the common
petiole. They do not always do this. Divided leaves, such as
those of Fig. 198 and 202, though ranked among the simple
sorts, are compound in the sense of having distinct blades,
but without articulation. Some of these blades are apt to be
confluent ; that is, a divided leaf is often in part merely parted,
as in the upper portion of Fig. 198. Such leaves are so inter-
mediate between simple and compound that it becomes indiffer-
ent, or a matter of convenience to be settled by analogy, under
which head or by what language they shall bo described. How-
ever, most leaves are so constituted as to leave no doubt whether
they are simple or compound.
194. The leaflets of a compound leaf being homologous with
the lobes or segments of a simple leaf, indeed being such segments
fully isolated, the two sorts fall under the same types. A pin-
nately veined simple leaf is the homologue of one kind of com-
pound leaf; a radiately veined leaf, of the other. That is,
compound leaves are either pinnate or palmate.
195. Pinnate Leaves (Fig. 203-205) are those in which the
leaflets are arranged along the sides of a petiole, or rather of its
THEIR STRUCTURE AND FORMS.
101
prolongation, the RHACHIS, which answers to the midrib of a pin-
nately veined simple leaf. There are three principal sorts, and
some subordinate ones. That is, a pinnate leaf ma}- be
Impari-pinnate, or pinnate with an odd leaflet, i. e. a terminal
one, as in Fig. 203 ; and this is the commoner case.
Cirrhiferous Pinnate, or pinnate with a tendril (Fig. 204) , as
in the proper Pea tribe and Bignonia. Here either the termi-
nal leaflet only, or the upper lateral leaflets also, are replaced
by tendrils.
Pari-pinnate, or Abruptly Pinnate, destitute of a terminal leaflet
or of an}' thing answering to it, as in Fig. 205.
Interruptedly Pinnate denotes merely a striking inequalit}' of
size among the leaflets : Lyrately Pinnate, one in which the termi-
nal leaflet is largest and the lower small.
196. Palmate or Digitate Leaves (Fig. 206, 93) are those in
which the leaflets all stand on the
summit of the petiole. Digitate
(fingered) was the old name, when
the term palmate was restricted to
a simple but palmate!}' lobed leaf
of this t}'pe. But since the time
of DeCandolle the two names have
been used interchangeably. Pal-
mate leaves have no primary dis-
tinction into sorts, except as to
the number of leaflets. These can 200
never be very numerous ; but there are fully a dozen in some
FIG. 203. An impari-pinnate or odd pinnate leaf. 204. Pinnate with a tendril
205. Abruptly pinnate leaf of a Cassia.
FIG. 206. Palruately or digitately 5-foliolate leaf of a Buckeye, JEseulus.
102 MORPHOLOGY OF LEAVES.
Lupines. More commonly there are only five to nine, or only
three, rarely two, or even a single one.
197. Number of leaflets may be indicated by an adjective
expression composed of the proper Latin numeral prefixed to
foliolale (Foliolum, diminutive of folium, answering to leaflet) .
Thus, bifoliolate, of two leaflets ; trifoliolate, of three leaflets ;
quadrifoliolate, of four ; quinquefoliolate, of five : plur if olio late, or
multifoliolate, of several or numerous leaflets, &c. These terms
are still more descriptive when accompanied b}' the word pin-
nately or palmately, indicative of the kind of compound leaf; as,
palmately or digitately trifoliolate (common Clover- leaf, Fig. 211),
or 5-foliolate, as in Buckeye (Fig. 206), and so on. Also, pinnately
\-foliolate, as in Fig. 205, or 17-foliolate, as in Fig. 203;
pinnately trifoliolate, as in Phaseolus, and in the low Hop-Clover,
Trifolium procumbens. 1
198. But, in either class of compound leaves, the leaflets may
be reduced to a minimum number. A pinnately trifoliolate leaf
is one of the impari-pinnate kind reduced to three leaflets, to one
pair and the odd one ; and this is distinguished from a palmately
trifoliolate leaf by the attachment of the pair at some distance
below the apex of the petiole, and by the articulation above this,
which marks the junction of the terminal leaflet's petiole (or its
base, if sessile) with the rhachis or common petiole.
199. Unifoliolate compound leaves (by no means a direct con-
tradiction in terms) are by this articulation distin-
guished from simple leaves which they simulate.
See the leaf of the common Barberry, Fig. 207.
In other species, of the Mahonia section, the leaves
are all pinnatety 3-9-foliolate, w r ith well-developed
common petiole : in the true Berberis, they are all
thus reduced to the terminal and long-petiolulate
leaflet, on an almost obsolete petiole. Orange and
Lemon leaves are in similar case, but with the joint
close to the blade. A comparison with near rela-
tives shows that these are also unifoliolate leaves
of the pinnate kind ; though this could not be ascertained by
inspection.
200. Decompound or Twice and Thrice Compound Leaves. These
are to once pinnate or once palmate leaves what the latter are to
1 In pinnate leaves, each leaflet usually has its opposite fellow, and the
number may be indicated by the pairs, as nnijugate, bijuyate, trijugate, and
plurijugate, according to the number oijuga, or pairs.
FIG. 207. Unifoliolate leaf of Berberis vulgaris, with partial petiole articulated to
the extremely short true petiole.
THEIR STRUCTURE AND FORMS. 103
simple leaves. As leaflets may be toothed, lobed, or parted, so
what answers to a single leaflet may appear as leaflets of a second,
or again of a third, or even of a fourth order. Decompound is a
good general name for all more than once compounded leaves ;
but the name has been applied rather to irregularly many-times
parted or dissected leaves (such as those of Dicentra) , or to
those more than thrice compounded. Of regularly twice or
thrice compound leaves, the commonest are the
Bipinnate or Twice Pinnate, of ordinary
occurrence in the Mimoseous and Caesalpi-
neous, but not in the Papilionaceous, Legu-
minosse. Fig. 208 represents a bipinnate leaf
of the Honey Locust (Glcditschia), with the
variation (common with that tree) that some
of the partial petioles, in this figure only the
lowest, bears a single leaflet, while the others
are extended into secondary rhachises fur-
nished with numer-
ous leaflets, mostly
in the abruptly pin-
nate st3 T le. On the
same tree, the earlier
loaves, which are
clustered on short
spurs, are simply
pinnate. The large J
loaves of Gymnocladus are similarly and abruptly bipinnate,
FIG. 208. A bipinnate and multifoliolate leaf of Gleditscliia or Honey Locust.
FIG. 209. Bipinnate leaves of Sensitive Plant, Mimosa pudica, with approximate
pinnse.
104
MORPHOLOGY OF LEAVES.
except at the base, which is simply pinnate or with one or two
pairs of simple leaflets.
Tripinnate or Thrice Pinnate leaves of a regular sort are rare ;
but, with some irregularity, they occur in many species, as in
Aralia, &c. This extent of division, and even much greater, is
common in Ferns.
Digitate- Pinnate is where the primary division of the petiole is
on the palmate or digitate plan ; the secondary, on the pinnate.
This seems to be the case in the Sensitive Plant, Mimosa pudica,
Fig. 209. But the leaf is here truly bipinnate with the primary
divisions very crowded at the apex of petiole.
Conjugate- Pinnate is the same arrangement, with the primary
divisions a single pair, at the apex of the petiole, and the leaflets
pinnately arranged on these.
Digitately or Palmately Decompound in a nearly regular way
is not an uncommon case.
Usually, the petiole is succes-
sively three-forked, as in Fig.
210, when the leaf is said to
be biternate (twice ternate),
triternate (thrice ternate), or
quadritemate (four times ter-
nate), &c., according to the
number of times it divides, or
2-3-4-times ternately compound.
210 The ultimate divisions in such
cases of threes are commonly of the pinnately trifoliolate type.
201. Pinnae is a convenient name for the partial petioles of a
bipinnate leaf, taken together with the leaflets that belong to
them. Thus, the Sensitive Plant, Fig. 209, has four pinnae, or
two pairs ; the Honey Locust, Fig. 208, a greater number.
When such leaves are still further compounded, the pinnae of
higher order, or the ultimate ones, take the diminutive term of
PINNULJE or PINNULES. The blades these bear are the Leaflets.
202. The Petiole or Leafstalk is a comparatively unessential
part of the leaf. It is often wanting (then the blade is sessile) ;
it may be absent even in compound leaves of the palmate type,
the leaflets rising side by side from the stem. When present, it
is usually either round, or half-cylindrical and channelled on the
upper side. In the Aspen, it is flattened at right angles with
the blade, so that the slightest breath of air puts the leaves in
motion. Sometimes it is much dilated and membranaceous at
FIG. 210. Quadri-ternately compound or ternately decompound leaf of Tlialictrum
Cornuti.
THEIR STRUCTURE AND FORMS.
105
base, as in many umbelliferous plants ; sometimes it forms a
sheath, occasionally it is bordered with appendages, &c. Peti-
oles may assume special functions, to be hereafter considered.
The woody and A^ascular tissue runs lengthwise through the
petiole, in the form usually of a definite number of parallel
threads, to be ramified in the blade. The ends of these threads
are apparent on the base of the leafstalk when it falls off, and on
the scar left on the stem, as so many round dots (Fig. 81, 85,
91), of a uniform number and arrangement in each species.
203. Partial Petioles are the divisions of the petiole in a
compound leaf. The footstalk of a leaflet takes the diminutive
name of PETIOLULE.
204. Stipules (157) are lateral appendages, one each side of
the base of the petiole, sometimes free
from it and from each other (Fig.
142), sometimes attached by one
edge to its base (Fig. 211), some-
times united with each other into a
single body (Fig. 212) in various
ways or degrees. In the latter case,
they usually appear to be within the
base of the leaf or leafstalk ; or, as
in the Plane-tree, they may be joined
into one over against the leaf, as
if opposite to it, but their normal
position is supposed to be lateral or
marginal to the petiole. Sometimes
they are foliaceous in appearance and
in function ; sometimes they are dry
and colorless or scale-like, reduced to
mere epidermal tissue, and evidently
functionless ; sometimes (as in Mag-
nolia, Fig. 81, Fig-tree, and Beech),
they serve as bud-scales, and fall when
the leaves develop ; sometimes the}' are
reduced to a mere bristle, or take the 2 n 212
form of a spine, as in the Locust (Robinia). Between salient
expansions or wing-like margins of tli3 base of the petiole,
such as those of the Saxifrage tribe, and stipules adnate to the
margins of the petiole, as in most Rosacea?, there is no clear
limitation. But presence or absence of stipules generally runs
FIG. 2H. Clover-leaf, with adnate stipules. 212. Odireate stipules (ochrea or
of Polygonum orientale, sheathing the stem for some distance, and ending in a spread-
ing border.
106 MORPHOLOGY OF LEAVES.
through a natural order. Yet what are called stipules in one order
ma}' pass for expansions or appendages of the petiole in another.
In Spergularia, some stipules are connate around the base of the
pair of leaves, including them as well as the stem in the sheath. 1
205. Stipules, which are normally a pair, may unite into one
bodj', either adnate to the inner face of the leaf, as in some
species of Potamogeton, or united opposite the leaf, as in Plane-
tree, or united inter se in a sheath, as in Potygonum. Also when
the leaves are opposite and the stipules thus brought into prox-
imity, the adjacent half stipules of the two leaves may coalesce,
and present the appearance of only two stipules to two leaves,
as in many Rubiaceae. A notch or fork at the apex often
indicates the composition.
206. Sheathing stipules, like those of Polygonum (Fig. 212),
are said to be ochreate, or (better) ocreate ; the sheath, thus
likened to a leggin or the leg of a boot, is an OCHRE A, as written
by Willdenow, or better OCREA.
207. The LIGULE of Grasses (Fig. 150) is seemingly a thin and
scarious extension of the lining to the sheath which answers to
petiole in such leaves : it projects at the junction of the sheath
and blade, there forming a kind of ocrea ; and it is generally
regarded as a sort of stipule.
208. Stipels (Stipella) are as it were stipules of leaflets, which
are common in certain tribes of Papilionaceous Legummosse, e. g.
in the Phaseoleae, in Wistaria, Locust, &c. ; also in Staplrylea.
The}^ are small and slender, and, unlike stipules, they are single
to each leaflet, except to the terminal one, which has a pair. As
leaves furnished with stipules are said to be stipulate, so leaflets
with stipels are stipe/late.
209. Some unusual modifications of leaves as foliage. In
leaves as illustrated thus far, it is the lamina or blade which is
expanded to do the work of foliage ; which is expanded hori-
zontally, so as to present upper and under surfaces, one to the
sky, the other to the ground ; which is bilaterally symmetrical
or substantially so, the two lateral halves being nearly if not
quite alike ; and which is affixed to the stem at the basal margin,
or some part of it, with or without a petiole. Various deviations
or apparent deviations from this pattern occur. Some of them are
of comparatively small account and simple explanation, such as
210. Inequilateral Leaves, being unsymmetrical by the much
greater development of one side. This is illustrated in the
whole genus Begonia (as in Fig. 161), consisting of many spe-
1 As pointed out by Prof. A. Dickson, in Nature, xviii. 507.
THEIR STRUCTURE AND FORMS.
107
cies, some of which are moderate!}', and most of them strikingly,
oblique in this way. Elm-leaves, and the like, are more or less
inequilateral at the base.
211. Connate and Perfoliate Leaves. These are explained by
the union of con-
tiguous leaf-edges.
Peltate leaves, to
which a paragraph
has already been
given (182), come
under the same
head ; the seeming
attachment of the
petiole to the lower
face of the blade
being the result of
a congenital union
of the edges of the
sinus. In a sessile
leaf, when such a 213 214
union takes place, it surrounds and encloses that portion of the
stem; which is thus perfoliate. (Fig.
213, 214.) It is the stem which is
literally perfoliate, i. e. which seem-
ingly passes through the leaf ; but it
is customary, though etymologically
absurd, to call this a perfoliate leaf!
Uvularia perfoliata (Fig. 213), in the
later growth of the season, reveals the
explanation of the perfoliation : the
base of the lower leaves conspicuously
surrounds and encloses the stem : that
of the upper is merely cordate and
clasping ; the uppermost simply ses-
sile by a rounded base. Baptisia
perfoliata (Fig. 214) is a more
strongly marked case of perfoliation.
But there are good morphological
reasons for inferring that this seemingl} 7 simple leaf consists of
a pair of stipules and a leaflet combined. An occasional mon-
strosity verifies this supposition.
FIG. 213. Leafy branch of Uvularia perfoliata.
FIG. 214. Leafy and flowering branch of Baptisia perfoliata.
FIG. 215. Lonicera flava, a wild Honeysuckle, connate-perfolinte as to the upper
leaves.
108 MORPHOLOGY OF LEAVES.
212. When leaves are opposite, the perfoliation (such as that
of Honeysuckles, Fig. 215) is obviously the result of a congeni-
tal union of the bases of the pair by their contiguous edges.
Leaves connate in this way by narrow bases are not rare nor
remarkable ; but when the two are thus coalescent into one broad
foliaceous body, giving this appearance of perfoliation, the term
connate-perfoliate is used to express it.
213. Vertical Leaves, those with blades of the ordinary kind,
but presenting their edges instead of their faces
to the earth and sky, or when erect with one
edge directed to the stem and the other away
from it, are not uncommon. They prevail in
the Australian Myrtaceas, &c., and occur with
less constancy in the Californian Manzanitas,
and in a great variety of herbs and shrubs. The
anomaly involves no exception to the rule that a
leaf-blade is always expanded in the horizontal
plane, when expanded at all ; for, except in equi-
tant leaves, it is the result of a twist of the petiole
or of the blade itself. 1 In strong^ marked
cases, or in most of them, the organization of
the epidermis and superficial parenchyma and
the distribution of the stomata are the same on
both faces.
214. Equitant Leaves are vertical on a different
plan. They are conduplicatc, i. e. are folded
2 1 7 2'.C
together lengthwise on their middle, the upper surface thus con-
cealed within, the outer alone presented to the air and light.
1 Silphlum laciniatum, the so-called Compass Plant, and (hardly less so)
S. terebinthinaceum, are good instances of the kind, most of the leaves
making a half-twist, the radical ones by their long petioles. In the former
species, the pinnately parted blade occasionally makes a farther twist, so as
to bring the upper part into a plane at right angles to the lower. The
blades place themselves in various directions as respects the cardinal points ;
but on the prairies the greater number affect a north and south direction of
their edges, a peculiarity first pointed out, in the year 1842, by General
B. Alvord, U. S. A.
FIG. 216. Equitant erect leaves of Trig, with the rootstopk.
FIG 217. A section across these leaves at the base, showing the equitant charnpte?.
THEIR STRUCTURE AND FORMS.
109
Being two-ranked and closely crowded, the outer ones at their
base fold over or bestride the inner (as shown in the sectional
diagram, Fig. 217), whence the name of equitant. Above, the
contiguous halves of the inner face congenially cohere, and so
produce the sword-shaped or linear vertical blade which is
characteristic of Iris (Fig. 216) and the Iris family. In most
there is a farther complication, of an excep-
tional kind, viz. the development backwards
of a portion of blade from the midrib, often
forming most of the upper part of such leaves,
which therefore may really be said to develop
in the vertical plane.
215. Leaves with no distinction of Parts, i. e.
of blade and petiole. This is the case in Iris
(Fig. 21G), Daffodil, the Onion, and perhaps
of most parallel-veined leaves of Endogens.
Those expanded in the horizontal plane may
however be regarded as sessile blades : those
which are not expanded, but filiform, or needle-
shaped (acicular), or awl-shaped (subulate), may
be regarded either as homologous with petioles,
or as unexpanded blades, which amounts nearly
to the same thing where there is no trace of a petiole at base.
Under this head may be ranked the leaves of Pines (Fig. 248) ;
also both the subulate and the
scale-shaped and adnate leaves
of Arbor Vitae, Red Cedar (Jiuri-
perus Virginiana) , and other trees
of the Cypress tribe. (Fig. 218.)
216. Stipules serving for Blade.
Lathyrus Aphaca is a good in-
stance of this (Fig. 219) ; the
petiole becoming a tendril, the
leaflets which its relatives bear
being wholly wanting, the ample
foliaceous stipules assume the
appearance of leaves. In some 219
other species of Lathyrus, and in the Pea, equally large stipules
share with the pair or pairs of leaflets in the functions of foliage.
On morphological evidence, we judge that the singular leaves of
FIG. 218. A twig of Arbor Vitae, with both awl-shaped and scale-shaped leaves.
FIG. 219. Lathyrus Aphaca: portion of stem, bearing a single leaf, which consists
of a pair of foliaceous stipules, and a petiole in the form of a tendril; in its axil a
flower-stalk.
110 MORPHOLOGY OF LEAVES.
Baptisia perfoliala, shown in Fig. 214, are not simple blades,
but each a pair of stipules, with or without a terminal leaflet, all
completely confluent into one body. The related species of the
genus have trifoliolate leaves and foliaceous stipules ; hence these
simple leaves without stipules are best explained in this wa}*.
217. Phyllodia, or Petioles serving for Blade. Sometimes the
petiole develops foliaceous margins, or wings, as in the Bitter
Orange and in Rhus copallina. These are efficient as foliage in
proportion to their size. These are not to be confounded with the
case in which a petiole specially develops as a blade-like organ,
which usurps the office of foliage. A petiole-blade of this kind
is named a PHYLLODIUM. Occurring only in Exogens, phyllodia
are generally distinguished from true blades by the parallel
venation, and alwa} r s by their normally vertical dilatation ; i. e.
the}', without a twist, present their edges instead of their faces to
the earth and sky. The common and most familiar phyllodia are
those of Acacias in Australia (Fig. 223, 224), where they form
the adult foliage of over 270 out of less than 300 species. The
true lamina of these is bipinnate. It appears on seedlings, and
occasionally on later growths. Several South' American species of
Oxalis produce phyllodia. So likewise do our tubular or trumpet-
leaved species of Sarracenia in that portion of the foliage which
develops the pitcher imperfectly, or not at all. Indeed, all
Sarracenia-leaves are phyllodia with the back in most of them
hollowed out into a tube or pitcher ; and the terminal hood
answers to the blade.
3. LEAVES SERVING SPECIAL OFFICES.
218. Leaves ma} T serve at the same time both their ordinaiy
and some special use, or even more than one special use. For
example, in Nepenthes (Fig. 222) there is a well-developed
blade, usually sessile, which serves for foliage, a prolongation of
its tip into a tendril, which serves for climbing, then an extraor-
dinary dilatation and hollowing of the apex of this into a pitcher
for a very special use, and a peculiar development of the apex
of this into a lid, closing the orifice during growth. Among the
special purposes which leaves subserve, and the study of which
connects singularities of morphology with teleology, the most
remarkable is that of
219. Leaves specialized for the Utilization of Animal Matter.
This occurs in leaves which also assimilate, or do the ordinary
work of vegetation ; and the special function is usually taken up
by some particular portion of the organ. The details of this
LEAVES SERVING SPECIAL OFFICES.
Ill
subject which has of late become highly interesting belong
to physiology, and therefore to the following volume, to which
all historical references are relegated. Only the morphology of
such leaves is here under consideration.
220. As Ascidia or Pitchers, vessels for maceration, &c. These
occur in several widely different
families of plants. The commonest
are those of the Sarracenias, natives
of Atlantic North America. They
are evident!}' plryllodia (217), the
cavity being a hollowed dorsal por-
tion : the wing-like or foliaceous por-
tion, alwa}"S conspicuous and forming
the ventral border, makes the whole
organ or most of it in the earlier
leaves of the tubular species. The
pitchers of S. purpurea (Fig. 221,
225), the only species which extends
north of Virginia, are open cups,
half filled with water, much of which
may be rain, in which abundance of
insects are usually undergoing macer- 22.5"
ation. In 8. variolaris (Fig. 226), the hooded summit, answer-
ing to the blade of the leaf, arches over the mouth in such wise
FIG 220. Pitchers of Heliamphora; 221. of Sarracenia purpurea ; 222. of Nepenthes.
223. A phyllodium of a New Holland Acacia. 224. The same, bearing a reduced com-
pound blade.
FIG. 225. Pitcher-leaves of Sarracenia purpurea ; one of them with the upper part
cut away.
112 MORPHOLOGY OF LEAVES.
as to mostly exclude the rain ; in S. psittacina (Fig. 227) the
inflexed and inflated hood completely excludes it. The water
which these contain is undoubtedly a secretion.
All entrap flies, ants, and various insects, which
in most species are lured into the pitcher 03- a
sweetish secretion around or at some part of the
orifice. 1 Few that have entered ever escape ;
most are decomposed at the bottom of the cavity.
In Darlingtonia Calilbrnica (Fig. 228) , the Cali-
fornian representative of Sarracenia, the inflated
hood guards against all access of rain, while the
orifice is freely open to ftying insects from be-
neath ; and a singular two-forked appendage, like
to a fish-tail (probably the homologue of the
blade) , overhangs the front. The inner face of
this appendage is besmeared with the sweet and
viscid secretion which allures insects to the open-
ing. In this and in Sarracenia variolaris, the
sweet secretion in the early season is continued
upon the edge of the wing, forming a saccharine
trail which leads from near the ground up to the
orifice of the pitcher. 2 Fig. 220 represents pitchers
of Heliamphora, a little-known South American
representative of Sarracenia. Its wing is narrow
and inconspicuous, the mouth widely open and
directed upward, and the hood reduced to a
minute and upright, probably functionless ap-
pendage. In Cephalotus an anomalous plant
of Australia, of uncertain affi-
mt} T the leaves for foliage
are dilated phyllodia ; among
them are others completely
transformed into stalked and
short pitchers, with thickened JK^ w
rim and a well-fitting lid,
hinged by one edge, Fig. 229.
The particular morphology of the parts is not well made out.
1 This sweet secretion, which at times is very obvious in the southern
species, has also been detected by Mr. Edward Burgess in S. purpurea; but
it is rarely seen, and probably plays no important part in the capture and
drowning of the multitude of insects which these pitchers are apt to contain.
2 This trail was discovered by Dr. J. H. Mellichamp, of South Carolina.
See Proc. Am. Association for Advancement of Science, xxiii. 113 (1871).
FIG. 226. Pitcher of Sarracenia variolaris. 227. Same of S. psittacina.
LEAVES SERVING SPECIAL OFFICES. 113
221. The pitcher-bearing leaf in Nepenthes has been referred
to (218, Fig. 222) : of this there are various species, all of them
somewhat wood}' climbing
plants of tropical Asiatic
and African islands of
the southern hemisphere,
some of them familiar in
conservatory cultivation.
Here the tendril may be
regarded as a prolonged
extension of the midrib
of the blade, and the
pitcher, with its hinged
lid, as a peculiar development from its apex. The
water contained in the pitcher is a secretion, much
of which appears before the lid opens ; and a sweetish
excretion at the orifice lures insects. The presence
of these in the pitcher increases the watery secre-
tion in which the animals are drowned ; and this
secretion is ascertained to have a certain digestive
power. 1
222. The aquatic sacs of Utricularia or Bladder-
wort are diminutive ascidia, always under water,
and with lid opening inward, like a valve, preventing the exit
of minute animals entrapped therein. 2 Morphologically, the}'
are doubtless leaves or parts of leaves.
223. As Sensitive Fly-traps. The leaves of all species of
Drosera or Sundew are beset with stout bristles tipped with a
gland, which secretes and when in good condition is covered
b} r a drop of a transparent and very glairy liquid, sufficiently
tenacious to hold fast a fly or other small insect. Adjacent
bristles, even if not touched, in a short time bend towards those
upon which the insect rests, and thus bring their glands also
into contact with it. In Drosera filiformis, the leaves are fili-
form, with no distinction of petiole and blade. In D. rotundifolia
and other common round-leaved species, there is a clear distinc-
1 This was first made out by J. D. Hooker, and announced in his address,
as President of the British Association for the Advancement of Science, at
Edinburgh, 1871.
2 Darwin, Insectivorous Plants, 305. Colin, Beitrage zur Biologie der
Pflanzen, 1875. Mrs. Treat, in The Tribune, New York, September, 1874,
and Card. Chron. 1875, 303.
FIG. 228. Pitcher of Darlingtonia Californica. 229. Pitcher of Ceplialotus follicu-
laria, with lid open.
114 MORPHOLOGY OF LEAVES.
tion of petiole and blade, and the stalked glands thickly beset
the whole upper surface of the latter. A small insect alighting
thereon is helpless, and is soon touched by all the glands within
reaching distance ; also the blade itself commonly incurves,
taking part in the general movement. It has recently been
demonstrated that the captured insect is fed upon, and that the
plant thereby receives nourishment. Here leaves which do the
normal assimilative work of vegetation, but somewhat feebly
(having a comparatively small amount of chloroplryll) , have
also the power and the habit of obtaining readj'-organized food
by capture, and are benefited b} T it.
224. Species of Drosera inhabit most parts of the world, and
the genus is numerous in species. A near relative, Dionsea, is
of a single species, D. muscipula (Venus's Fty-trap), inhabiting
only a limited district in the sandy eastern border of North
Carolina. It is more strikingly sensitive and equally carnivo-
rous, but in a different way. It is destitute of stalked and viscid
glands. The apparatus for capture and digestion is the two-
valved body at the top of each leaf. (Fig. 230, 231.) If this
FIG. 230. A plant of Dionsea mnscipula, reduced in size. 231. Three of the leaves,
of almost the natural size ; one of them open, the others closed. Probably a fly is never
caught by the teeth, in the manner here represented.
LEAVES SERVING SPECIAL OFFICES.
115
be taken for the leaf-blade, the part below would be a broadly-
winged foliaceous petiole. If the latter be the true blade, the
apparatus in question must be reckoned as a peculiar terminal
appendage. Both are moderately green, and act as foliage.
The specially endowed terminal portion acts also in a decidedly
animal-like manner. When either of the three or four slender
bristles of the upper surface are touched, the trap suddenly
closes, by a movement ordinarily quick enough to enclose and
retain a fly or other small insect. The intercrossing of the stout
marginal bristles detains the captive, unless it happens to be
small enough to escape by the intervening little openings.
Otherwise, the sides soon flatten and are brought firmly into
contact, and a glairy secretion is poured out from numerous
immersed glands : this, with the extracted juices of the macerated
insect, is after some time reabsorbed ; the trap, if in a health}'
condition, now re-opens and is read}' for another capture. For
references to the now copious literature of this whole subject, and
for its physiological treatment, the succeeding volume should be
consulted.
225. Leaves for Storage. Nutritive matter is stored in leaves
in many cases, and not rarely in leaves which at the
same time are subserving the purpose of foliage.
This occurs in all fleshy leaves, to a greater or less
extent, according to the degree of thickening or
accumulation. The leaves of the Century Plant
or Agave, for instance, are green and foliaceotisly
efficient at the surface, while the whole interior is
a store-house of farinaceous and other nutritious
matter, as much so as is a potato. The leaves
of various species of Aloe, Mcseinbryanthemum,
Sedum, and other "succulent" plants (in which
a large part of the accumulation is water) arc not
rarely so obese as to lose or much disguise the
foliaceous appearance. Sometimes one portion of
a leaf is of normal texture and use, whib another
is used as a reservoir for the nourishment which the
foliaceous part has produced. Fig. 232, a loaf from
the bulb of White Lily, the base of which forms
one of the bulb-scales, is an instance of the kind. 1 The most
1 In Dicentra Cucullaria and (more strikingly from the sparseness of the
grains) in I). Oanadensis, the matter elaborated in the much dissected blade is
conveyed to the very base of the long petiole, and there deposited in a con-
Fid. 2H2. A radical leaf of Tie White Lily, with its base thickened into a bulb-scale,
which id cut across to show its thickness.
116
MORPHOLOGY OF LEAVES.
decisive instance of leaves used for storage of food is in that
material provision for the nourishment of the embryo in germi-
nation, in which the first leaves, the cotyledons, are turned to this
account. (21-37, &c.) After or
while discharging this special duty,
the cotyledons may fulfil their gen-
eral office, by serving as foliage (as
in Maples, Fig. 8, and Pumpkins,
Fig. 47) ; or, through various inter-
mediate conditions, the} r may be
wholly devoted to storage, as in
'the Pea, Oak, Horsechcstnut, &c.
(Fig. 37-43.)
226. Leaves as Bulb-scales, how-
ever, are for the most part wholly
applied to this use, being leaves
reduced to short scales or to
concentric coats, and thickened
throughout by nutritive deposit.
The accumulation of such leaves
forms the mass of the bulb, as of
the Lily, Fig. 118, Onion, Fig.
113, &c., also of bulblets. (120.)
227. Leaves as Bud-scales, being
for protection of nascent parts, have
been explained under buds. (70.)
The evidence of foliar nature af-
forded by transition is well exhib-
ited by the Sweet Buckeye, although
the whole series of gradations, from
bud-scales to compound leaves,
is seldom seen united in one bud,
as in Fig. 233. In this case, the
bud-scales are homologous with
petioles. In Magnolia, they consist
of stipules (Fig. 81, 82) : in the
Lilac, they arc homologous with leaf-blades. The two pairs of
bud-scales which subtend .and protect through winter the nascent
head of flowers of Cornus florida are morphologically the apex of
centrated condition, in the form of a solid grain, which remains for next
year's use, the whole leaf except this thickened base dying away at the close
of the short season's growth.
FIG 233. Leaves of a dcvolopinsr bud of the Low Sweet Buckeye (vEscnlns pnrvi-
flora), showing a nearly complete set of gradations from a scale to a compound leaf ol
five leaflets.
LEAVES SERVING SPECIAL OFFICES. 117
blades. When the blossoms develop in spring, these scales grow
from beneath, greatly expand, and become obovate or obcordate
petaloid leaves, the brown terminal notch
of which is the bud-scale, which was un-
able to take part in the vernal growth.
22 7 a . Leaves as Spines. All gradations
ma}' be found between spiny-toothed leaves
(as in Holly), in which teeth arc pointed and
indurated, and leaves which are completely
contracted into a simple or multiple spine.
Indeed, such a transition is seen in the Bar-
berry, Fig. 234. The foliar nature of such
spines is manifest from their position, sub-
tending a bud from which the foliage of the
season proceeds, and themselves not sub-
tended by any organ. In some Astragali,
the petiole of a pinnate leaf indurates into
a slender spine and persists, the leaflets
early falling. The spine in Fouquiera is a
portion of the lower side of the petiole or
midrib, indurated and persistent, the rest
of the leaf separating by splitting when it
has served its office. 1
228. Leaves adapted to Climbing. Some plants climb by the
action of the stem or of certain branches specialty adapted to
this purpose (99) : others gain the needful
support by means of their leaves (101) ; some-
times by an incurvation of the tips, either of
a simple blade as in Gloriosa, or small partial
blades, as in Adlumia, and often in Clematis,
thereb}' grappling the support ; sometimes by
the petiole making a turn or two around a
support (as in Maurandia, climbing Antirrhi-
nums, Rhodochiton, and Solanum jasminoides,
Fig. 235) ; sometimes b}' the transformation
of one or more leaflets of a compound leaf
into tendrils, as in the Pea and Vetch (Fig.
204) ; sometimes by the suppression of all the
leaflets and the conversion of the whole petiole into a tendril, as
in Lathyrus Aphaca (Fig. 219) ; and perhaps by the conversion
1 Described in Plants Wrightiaiue, ii. 63.
FIG. 234. A vernal shoot of common Barberry, showing a lower leaf in the normal
stare; the next partially, those still higher completely, transformed into spines.
FIG. 235. Solanum jasminoides, climbing by coiling and at length indurating petioles.
118 MORPHOLOGY OF LEAVES.
of a pair of stipules into tendrils in Smilax. At least the ten-
drils here occupy the position of adnate stipules. The tendrils
of Cucurbitaceae are peculiar and ambiguous, on account of their
lateral and extra-axillary position and the manner in which the
compound ones develop their branches. But they are doubtless
partly if not wholly foliar. 1
229. Petaloid Leaves, Bracts. Certain leaves, situated near to
flowers, and developing little or no chlorophyll in their paren-
chyma, exchange the ordinary green hue and herbaceous texture
for the brighter colors and more delicate structure which are
commonly seen in and thought to characterize flower-leaves.
Such are said to be colored, meaning, as applied to foliage, of
some other color than green. As petals are the type of such
colored parts, they are said to be petaloid, i.e. petal-like. They
are like petals, moreover, in one of the purposes which these sub-
serve. (299.) Examples of these petaloid leaves are seen in the
shrubby Mexican Euphorbia called Poinsettia, in Salvia spleu-
dens, most species of Castilleia or Painted Cup, also in the
white hood of Calla and Richardia ^Ethiopica (called Calla Lily),
and in the four white leaves which subtend the flower-head of
Cornus florida, and of the low herbaceous Cornel, C. Canaden-
sis. (Fig. 294.) Such leaves, being in proximity to flowers, and
all others which are within a flower-cluster or are borne by
flower-stalks, receive the special name of BRACTS. More usually
bracts are not petaloid, but different in size or shape from ordi-
nary leaves, either by abrupt change or gradual transition. Not
uncommonly they are reduced to scales or mere rudiments or
vestiges of leaves, of no functional importance.
230. Flower- Leaves. The morphology of leaves extends not
only to " the leaves of the blossom," more or less accounted as
such in common parlance, but also to its peculiar and essential
organs, the relation of which to leaves is more recondite. Their
morphology needs to be treated separately, and to be preceded
by a study of the arrangement of leaves and of blossoms.
1 The most satisfactory interpretation may be that of Braun and Wytller,
adopted by Eichler (Bliithendiagramme, i. 304) : that the flower of Cucur-
bita and its peduncle represent the axillary branch, the tendril by its side
answers to one of the bractlets (that of the other side being suppressed),
and the supernumerary branch springs from the axil of the tendril. This
makes of the tendril a simple leaf, of which the branches are the ribs. But
the tendril-divisions are evidently developed in spiral order, and in vigorous
growths occupy different heights on the tendril-axis. This favors Naudin's
view, that the main tendril is cauline, and its divisions leaves.
PHYLLOTAXY, OK LEAF- ARRANGEMENT.
119
CHAPTER IV.
PHYLLOTAXY, OR LEAF-ARRANGEMENT.
SECTION I. THE DISTRIBUTION OF LEAVES ON THE STEM.
231. PHYLLOTAXY (or Phyllotaxis) is the stud}' of the distri-
bution of leaves upon the stem and of the laws which govern it.
The general conclusion reached is, that leaves
are distributed in a manner to economize space
and have a good exposure to light, &c., and
that this economy on the whole results from
the formation of leaves in the bud over the
widest intervals between the
leaves next below. 1 Leaves
are arranged in a consider-
able variety of ways, which
all fall under two modes, the
Verticillnte and the Alter-
nate (13), but which may
also be termed the Cyclical
and the Spiral.
232. Alternate leaves are
those which stand singly,
one after another ; that is,
with one leaf to each node
or borne
on one height of stem. Verticillate leaves are
those with two or more at the same height of
stem, circularly encompassing it, i. e. forming
a Verticil or Whorl. Verticillate and whorled
are synonymous terms to denote this arrange-
ment. These two kinds of leaf-arrangement
are commonly ranked as three, viz. alternate,
opposite, and whorled. But the opposite is
onl}' the simplest case of the whorled, being
1 For the most comprehensive discussion of phyllotaxy in connection with
development, and in view of these relations, see Hofmeister, Allgemeine
Morphologic, 11, and Chauncey Wright, Mem. Amer. Academy, ix. 389.
FIG. 236, Alternate, 237, Opposite, 238, Verticillate or whorled leaves.
120 PHYLLOTAXY, OR LEAF-ARRANGEMENT.
that in which the members are reduced to two. This case is
so much commoner than whorls of three and of higher numbers
that it took from the first its special name of opposite, so that in
descriptions the phrase " leaves verticillate " implies more than
two leaves in the whorl. But it should be kept in mind that
"leaves opposite " is the same as u leaves in whorls of two."
233. The greater number of phomogamous plants (all but the
monocotyledonous class) begin with verticillate leaves, most!}*
of the simplest kind (t. e. cotyledons opposite) : some continue
verticillate throughout ; some change in the first leaves of the
plumule or after the first pair into alternate, and again into
verticillate in or toward the blossom, in the interior of which the
alternate arrangement may be again resumed. As Nature passes
'-ladily from the one mode to the other on the same axis, we
may expect that the two ma3' be comprised under some common
expression. But the}- have not yet been combined, except by
gratuitous or somewhat forced hypotheses ; so that for the
present they should be treated in morphology as primarily dis-
tuiet arrangements. 1
234. Verticillate or Cyclical Arrangement. Here the leaves
occup t y a succession of circles, or form whorls around the stem,
two, three, four, five, &c., in each whorl. According to the
number, the leaves are opposite, ternate, qnaternate, (juinate, and
so on. The characteristic of the individual whorl is that the
members stand as far apart from each other as their number
renders possible, i. e. they divide the circle equal!}'. Thus, when
only two, or opposite, their midribs or axes of insertion have an
angular divergence (as it is termed) of 180; when three, of
120 ; when four, 90 ; when five, 72.
235. The characteristic of the whorls in relation to each other
is, that the members of successive whorls stand over or under
the intervals of the adjacent ones. In other words, successive
whorls alternate or decussate. This economizes space and light,
or gives the best distribution which the cyclical system is capa-
ble of. And it is in accordance with the general conclusion of
Hofmeister's investigation of the origin of phyllotaxic arrange-
ments in the nascent bud, viz. that new members originate just
over the widest intervals between their predecessors next below.
Thus, in opposite leaves or whorls of two (Fig. 237), the suc-
cessive pairs decussate or cross at right angles, and so four
1 It is readily seen that whorls may be produced by the non-development
of the internodes between the leaves of a series of two, three, five, or more
in alternate order. The difficulty is that the members of the next whorl do
not follow the rrder that they should upon this supposition.
DISTRIBUTION OF LEAVES ON THE STEM. 1-1
straight equidistant vertical ranks are produced. In ternate or
trimerous whorls there are six vertical ranks ; in quaternate or
tetramerous whorls, eight vertical ranks, and so on. 1
236. The cases in which successive
pairs of leaves do not decussate at right
angles, or the members of whorls are not
exactly superposed to intervals, but as it
were wind spirally (as in Dipsacus, man}'
Caryophyllaceae, &c.) , may some of them
be explained by torsion of the stem,
such as is veiy manifest in numerous in-
stances ; and others may be resolved into 239
instances of alternate leaves simulating or passing into whorls
by the non-development of internodes. 2
237. Alternate or Spiral Arrangement. Here the leaves are
distributed singly at different heights of the stem, and at equal
intervals as respects angular divergence. (Fig. 236.) This angu-
lar divergence (i.e. the angular distance of an}' two successive
leaves) differs in the various kinds of this system of phyllotaxy,
but is always large enough to place the leaves which immediately
1 These vertical ranks have, by some German botanists, been named
Orthosli clues ; but this technical Greek is no clearer and no shorter than the
equivalent English, which answers every purpose.
2 In Lilium Canadense, superbum, &c., with whorls of variable number
of leaves and vague relation to each other (when of the same number some-
times the members superposed), and above and below passing into the alter-
nate arrangement normal to the family, these whorls are evidently formed
of alternate leaves brought together by non-development of internodes.
Here may also be mentioned the not uncommon anomaly in Fir-cones,
notably those of Norway Spruce, the normal phyllotaxy of which is simply
spiral, but in occasional instances the cone is composed of pairs of opposite
scales, spirally arranged, /. e. the pairs not decussating at right angles, thus
forming double spirals. In the abnormal spruce-cones, the fractions usually
observed are $ or ^g, or, as expressed by Braun, (i)y 5 ^ and (%)-$
Braun's mode of notation for the ordinary succession (i. e. the decussation)
of opposite leaves is (|)i, the \ meaning that the two leaves of the pair are
half the circumference of the circle apart, the ^ denoting that each leaf of
the succeeding pair diverges one fourth of the circumference from the pre-
ceding. Braun finds cases in which pairs (and equally whorls) are super-
posed (e.g. certain species of Mesembryanthemum and Euphorbia), these
are expressed in this notation by the formula (), that is, the corresponding
leaves of the succeeding pair diverge 180 from their predecessors. He
recognizes alsn some cases of intermediate divergence; such as ()f in the
upper leaves of Mercurialis perennis, (%)$$ on certain stems of Linaria vul-
garis, (|)y 5 & exceptionally in the leaves of Epilobium angustifolium and the
scales of Norway Spruce, (i)-j 8 ^ exceptionally in the scales of Norway
Spruce. See Ordnung des Schuppen an der Tannenzapfen, 376, &c.
FIG. 239. Ground-plan diagram of six trimerous wborls, showing their alternation.
122 PHYLLOTAXY, OK LEAF-ARRANGEMENT.
follow each other in the ascending order upon different sides of
the axis : it also secures an advantageous spacing of the leaves
over the whole length of the axis. Their vertical distance from
each other of course depends on the length which the internodes
attain, which is a matter of growth and is very variable ; but
their angular distance is fixed in the kind or numerical plan of
the particular phyllotaxy, and is uniform throughout.
238. The leaves are said to be alternate, because they come
one after another, now on this side, then on that, as they ascend
the stem. The arrangement is said to be spiral, because if a line
be drawn or a thread extended from the base or insertion of one
leaf to that of the next higher, and so on, taking in all the leaves,
it forms a helix, more or less loose or close according to the
development of the internodes. (See Fig. 242.) This imagined
spiral line ascends continuous!}', without a break ; and on it the
leaves are equably laid down. 1
239. Almost all the ordinary instances of spiral plryllotaxy
belong to one series, having very simple arithmetical relations.
So that this may be taken as the type, and the few others re-
garded as exceptions or sometimes as modifications of it. The
kinds are simply designated by the number of vertical ranks of
leaves : the}' are technically named by prefixing the proper
Greek numeral to the word meaning row or rank. The arrange-
ment called
Distichous, or Two-ranked, is the simplest and among the com-
ynonest, occurring, as it does, in all Grasses and many other
/nonocotyledonous plants, in Lindens, Elms, and many dico-
tyledonous genera. Here the leaves are disposed alternately on
exactly opposite sides of the stem (as in Fig. 1 ) ; the second
leaf being the farthest possible from the first, as is the third from
the second ; the third therefore over the first, and the fourth over
the second, and so on, thus forming two vertical ranks. The
angular divergence is here half the circumference, or 180 ; and
the phyllotaxy may be represented by the fraction J, which desig-
nates the angular divergence, while its denominator expresses
the number of vertical ranks formed.
Tristichous, or Three-ranked, is the next in the series, and is
1 But when we reach a leaf which stands directly over a lower and older
one, we say that one set or spire is completed, and that this leaf is the first
of a succeeding set or spire. From analogy of such an open spire to the
closed cycle of a whorl of leaves, it is not unusual to designate the former
likewise as a cycle. Yet it is better (with Eichler) to restrict that term, and
the adjective cyclical, to verticillate phyllotaxy, or to whorls, to which it
properly and etymologically belongs.
DISTRIBUTION OF LEAVES ON THE STEM.
123
less common, though not rare in monocot}'ledonous plants. Fig.
240 illustrates it in a Sedge, and 241 is a diagram in horizontal
section, as of a bud ; both
extending to six leaves or
two turns of the spiral. The
fraction designates this
arrangement. - The angular
divergence, or distance o$
the axis of the first leaf
from the second, and so on,
is one third of the circum-
ference (or 120) : conse-
quently the fourth leaf comes
over the first, the fifth over
the second, the sixth over
the third, and so on ; that
is, the leaves fall into three
vertical ranks. The spiral
character here begins to be
manifest, or becomes so by
drawing a line on either fig-
ure from the axis or midrib
of the first leaf to that of
the second, and so on to the
sixth, forming a helix of
two turns. 1
Pentastichous, or Five-ranked, sometimes termed the guincuncial
arrangement. This is the most common in alternate-leaved
dicotyledonous plants. It is shown in Fig. 236 (on a branch
of Apple-tree), and by diagrams, displaying the spiral character,
in Fig. 242, 243. The angular distance from the first to the
second leaf (passing the shorter way) is of the circumference,
or 144. But the spiral line makes two turns round the stem,
on which six leaves are laid down, with angular divergence of f ,
1 The line is supposed to follow the nearest way, and the divergence is
counted as , this being the simplest and most convenient. If for any reason
the longer way is preferred, then the angular divergence would be expressed
by the fraction |.
FIG. 240. Piece of a stalk, with the sheathing bases of the leaves, of a Sedge-Grass
(Carex crns-corvi), showing the three-ranked arrangement. 241. Diagram of the cross-
section of the same. The leaves are numbered in succession.
FIG. 242. Diagram of position of six leaves in the five-ranked arrangement : a spiral
line is drawn ascending the stem and passing through the successive scars which mark
the position of the leaves from 1 to 6. It is made a dotted line where it passes on the
opposite side of the stem, and the scars 2 and 5, which fall on that side, are made
fainter. 243. A plane horizontal projection of the same; the dotted line passing from
the edge of the first leaf to the second, and so on to the fifth leaf, which completes the
turn ; as the sixth would come directly before, or within, the f
124
PHYLLOTAXY, OR LEAF-ARRANGEMENT.
and the sixth is the first to come over any one below : the
seventh comes over the second, the eighth over the third, &c.
The leaves are thus brought into five vertical ranks ; but these
five leaves are laid down on two turns of the
helix (the sixth beginning the second revolu-
tion) ; the angular divergence of the leaves in
order is f , or 144 ; the angular distance of
the vertical ranks, 72. This is a very advan-
tageous distribution for ordinary foliage on
erect or ascending branches. Its formula is , expressing the
angular divergence, the denominator also indicating the number
of vertical ranks, the numer-
ator indicating the number
of revolutions made in add-
ing one leaf to each rank.
Fig. 244 illustrates this ar-
rangement on a cone of
American Larch, the scales
of which are homologous
with leaves, the numbers
in sight are affixed, and those of the whole
cone displaj'ed on a plane at the side.
Octostichous, or Eight-ranked, a less common
arrangement, occurs in the Holly, Aconite,
the radical leaves of Plantago. It has the
angular divergence of 135, or | of the cir-
cumference, and the leaves in eight ranks,
the ninth over the first and at the completion
of the third revolution : it is therefore repre-
sented by the fraction |.
240. The obvious relations of the fractions
ii 3' f > t? representing the primary forms of
spiral phyllotaxy, are that the sum of any two
numerators is the numerator of the next suc-
ceeding fraction, and the same is true of the
denominators ; also the numerator is the same
as the denominator of the next but one pre-
ceeding fraction. Following these indications,
the series ma}' be extended to -f^, 5 8 T , ^ v |, , &c. Now these
FIG. 244. A cone of the small-fruited American Larch (Larix Americana), with
the scales numbered, exhibiting the five-ranked arrangement.
FIG. 245. An offset of the Houseleek, exhibiting the fi-13 arrangement; the leaves
in sight numbered, the 14th over the first, the 19th over the 6th, &c.
FIG. 246. Cone of White Pine (Pinus Strobus) with scales numbered from bottom,
and some secondary spirals marked.
DISTRIBUTION OF LEAVES ON THE STEM. 125
cases actually occur, and ordinarily only these. 1 The y\ and
^ T are not uncommon in foliage. The rosettes of the House-
leek exhibit the fy or thirteen-ranked arrangement, as also does
the cone of Pinus Strobus, the 14th leaf falling over the first.
(Fig. 246.) The ^ 8 T is perhaps little less common in foliage
upon very short internodes, as likewise are higher ranked
numbers ; and in many pine-cones and similar structures Jf
and plryllotaxy may be readily made out. This actual series,
, , f , |, &c., answers to and may be expressed by the con-
tinued fraction, \ + ,
t + i + i, &c. 2
1 When other instances are detected, they are found to belong to other
series, following the same law, such as the rare one of \, \, f, | 8 ^.
2 " The ultimate values of these continued fractions extended infinitely
are complements of each other, as their successive approximations are, and are
in effect the same fraction, namely, the irrational or incommeasurate inter-
val which is supposed to be the perfect form of the spiral arrangement.
This does, in fact, possess in a higher degree than any rational fraction the
property common to those which have been observed in nature; though
practically, or so far as observation can go, this higher degree is a mere
refinement of theory. For, as we shall find, the typical irrational inter-
val differs from that of the fraction f by almost exactly y^W' a quantity
much less than can be observed in the actual angles of leaf -arrangements."
" On this peculiar arithmetical property .... depends the geometrical one,
of the spiral arrangement, whijh it represents ; namely, that such an arrange-
ment would effect the most thorough and rapid distribution of the leaves
around the stem, each new or higher leaf falling over the angular space be-
tween the two older ones which are nearest in direction, so as to subdivide it in
the same ratio in which the first two, or any two successive ones, divide the
circumference. But, according to such an arrangement, no leaf would ever
fall exactly over any other; and, as I have said, we have no evidence, and
could have none, that this arrangement actually exists in nature. To realize
simply and purely the property of the most thorough distribution, the most
complete exposure of light and air around the stem, and the most ample
elbow-room, or space for expansion in the bud, is to realize a property that
exists separately only in abstraction, like a line without breadth. Neverthe-
less, practically, and so far as observation can go, we find that the fractions
| and T 5 ^, ^ 8 r , &c., which are all indistinguishable as measured values in the
plant, do actually realize this property with all needful accuracy. Thus,
| = 0.375, T \ = 0.385, and ? \ = 0.381, and differ from Tc [the ultimate value
to which the fractions of this series approximate, or what is supposed to be
the type-form of them] by 0.007, -f-0.003, and 0.001 respectively ; or they
all differ by inappreciable values from the quantity which might therefore be
made to stand for all of them. But, in putting k for all the values of the
series after the first three, it should be with the understanding that it is not
so employed in its capacity as the grand type, or source of the distributive
character which they have, in its capacity as an irrational fraction, but
simply as being indistinguishable practically from those rational ones."
Chaucey Wright, in Mem. Amer. Acad. ix. 387-390.
126 PHYLLOTAXY OR LEAF-ARRANGEMENT.
241 . The successive grades of angular divergence of alternate
leaves, as expressed in degrees, are
= 180 =144 A = 138 27' 41.54"
=:120 f 135 ^ = 137 8' 34.29"
and so on ; and be3*ond, if not in the latter cases, the differences
become quite too small for determination by inspection. They
all fall within the and as to amount of divergence : and they
form a series converging to a deduced typical angle of 137 30'
28", which, being irrational to the circumference, would place
no leaf exactly over any preceding one, but alternately and
more and more slightly on one and the other side of the vertical,
and so on, in an endless spiral. That is, according to Bravais,
the ranks in the higher grades tend to become curviserial, or*
actually become so; while in the lower grades they are obviously
rectiserial. Unless, indeed, there is some torsion of the axis,
by which the vertical ranks are rendered oblique, as is often the
case in cones of the Norwa}' Spruce. But, apart from this, the
difference between rectiserial in a high order and curviserial
soon becomes inappreciable. Any and all of the higher grades,
and practically one as low as the |, secures the utility of the
theoretical angle, viz., that " by which the leaves would be dis-
tributed most thoroughly and rapidly around the stem, exposed
most completely to light and air, and provided with the greatest
freedom for symmetrical expansion, together with a compact
arrangement in the bud." Even in the simpler grades of com-
monest occurrence, each leaf (according to Wright) is so placed
over the space between older leaves nearest in direction to it as
always to fall near the middle of the space, until the circuit is
completed, when the new leaf is placed over an old one. 1
242. It is to be noted that the distichous or \ varietj 7 gives
the maximum divergence, viz. 180, and that the tristichous or
^ gives the least, or 120 ; that of the pentastichous or f is nearly
the mean between the first two ; that of the f , nearly the mean
between the two preceding, &c. The disadvantage of the two-
ranked arrangement is that the leaves are soon superposed and
so overshadow each other. This is commonly obviated by the
length of the internodes, which is apt to be much greater in this
than in the more complex arrangements, therefore placing them
vertically farther apart ; or else, as in Elms, Beeches, and the
1 This corresponds with Hofmeister's general rule, that " new lateral
members have their origin above the widest gaps between the insertions of
the nearest older members." Yet the fact that the character of the leaf-
arrangement is laid down at the beginning in the bud does not go far in the
way of the mechanical explanation which he invokes.
DISTRIBUTION OF LEAVES ON THE STEM. 127
liko, the branchlcts take a horizontal position and the peti-
oles a quarter twist, which gives full exposure of the upper
fuce of all the leaves to the light. The and , with dimin-
ished divergence, increase the number of ranks ; the | and all
be}'ond, with mean divergence of successive leaves, effect a more
thorough distribution, but with less and less angular distance
between the vertical ranks.
242 a . The helix or primitive spiral upon which the leaves
successively originate ascends, sometimes from left to right,
sometimes from right to left, 1 commonly without change on the
same axis, and prevailingly uniform in the same species ; but
occasionally both directions occur in the same individual. The
earliest leaves of a stem or branch, or the last, are often on a
different order from the rest ; or (as already stated) the spiral
change into the cyclical, or vice versa.
243. The relation of the phyllotaxy of a branch to the leaf
from the axil of which the branch springs is somewhat various.
But in Dicotyledons, the first leaf or the first pair of the branch is
mostly transverse ; that is, the first leaves of the branch stand to
the right or left of the subtending leaf. In Monocotyledons, the
first branch-leaf is usually parallel to and facing the subtending
leaf, as shown in Fig. 304.
244. When the internodes are considerably lengthened, the
normal superposition of leaves is not rarely obscured by torsion
of the axis : indeed, this may equally occur in short internodes,
sometimes irregularly or in opposite directions, sometimes uni-
formly in one. Thus, in Pandanus utilis, or Screw-Pine, of
tristichous arrangement, the three compact vertical ranks be-
come strongly spiral by a continuous torsion of the axis. The
later leaves of Baptisia perfoliata, which are normally distichous,
become one-ranked by an alternate twist, right and left, of the
successive internodes.
245. When the internodes are short, so that the leaves approx-
imate or overlap, it is difficult or impossible to trace the suc-
cession of the leaves on the primitive spiral, but it is easy to
see which are superposed. The particular phyllotaxy may then
be determined by counting the vertical ranks, which gives the
denominator of the fraction. But in compact arrangements
these vertical ranks are commonly less manifest than certain
oblique ranks, which are seen to wind round the axis in oppo-
site directions. (See -Fig. 245, 246.) These are termed second-
ary spirals^ also by some parastichies. These oblique spiral
1 That is, of the observer and as seen from without. See p. 51, foot-note.
128 PHYLLOTAXY, OR LEAF-ARRANGEMENT.
ranks are a necessary consequence of the regular ascending
arrangement of parts with equal intervals over the circumference
of the axis ; and, if the leaves are numbered consecutively, their
numbers will necessarily stand in arithmetical progression on the
oblique ranks, and have certain obvious relations with the pri-
mary spiral which originates them, as will be seen by projecting
them on a vertical plane.
245". Take, for example, the f arrangement, where, as in the
diagram annexed to Fig. 244, the primitive spiral, written on a
plane surface, appears in the numbers, 1, 2, 3, 4, 5, 6, and so
on : the vertical ranks thus formed are necessarily the numbers
1-6-11; 4-9-14; 2-7-12; 5-10-15; and 3-8-13. But two
parallel oblique ranks are equally apparent, viz. 1-3-5, which,
if we coil the diagram, will be continued into 7-9-11-13-15 ; and
also the 2-4-6-8-10 continues into 12-14, and so on, if the axis
be prolonged. Here the circumference is occupied by two secon-
dary left-hand series, and we notice that the common difference
in the sequence of numbers is two ; that is, the number of the
parallel secondary spirals is the same as the common difference of
the numbers on the leaves that compose them. Again, there are
other parallel secondary spiral ranks, three in number, which
ascend to the right ; viz. 1-4-7, continued into 10-13 ; 3-6-9-12,
continued into 15 ; and 5-8-11-14, &c. ; where again the common
difference, 3, accords with the number of such ranks. This fixed
relation enables us to lay down the proper numbers on the leaves,
when they are too crowded for directly following their succes-
sion, and thus to ascertain the order of the primary spiral series
by noticing what numbers come to be superposed in the verti-
cal ranks. Thus, in the small cone of the American Larch
(Pig. 244), which usually completes only three heights of leaves,
the lowest, highest, and a middle one make a vertical row
which faces the observer. Marking this first scale 1, and count-
ing the parallel secondary spirals that wind to the left, we find
that two occup} T the whole circumference. From 1 , we number
on the scales of that spiral 3-5-7, and so on, adding the com-
mon difference 2, at each step. Again, counting from the base
the right-hand secondary spirals, we find three of them, and
therefore proceed to number the lowest one by adding this com-
mon difference, viz. 1-4-7-10 ; then, passing to the next, on
which the No. 3 has already been fixed, we carry on that se-
quence, 6-9, &c. ; and on the third, where No. 5 is already
fixed, we continue the numbering, 8-11, &c. This gives us in
the vertical rank to which No. 1 belongs the sequence 1-6-11,
showing that the phyllotaxy is of the five-ranked, or f order.
DISTRIBUTION OF LEAVES ON THE STEM. 129
It is further noticeable that the smaller number of parallel sec-
ondary spirals, 2, agrees with the numerator of the fraction in
this the f arrangement ; and that this number, added to that of
the parallel secondary spirals which wind in the opposite direction,
viz. 3, gives the denominator of the fraction. This holds good
throughout ; so that we have only to count the number of par-
allel secondary spirals in the two directions, and assume the
smaller number as the numerator, and the sum of this and the
larger number as the denominator, of the fraction which ex-
presses the angular divergence sought. For this, we must, how-
ever, take the order of secondary spirals nearest the vertical
rank in each direction, when there are more than two, as in all
the higher forms. But, in all, it is necessary to count only the
most manifest secondary spiral of each direction in order to
lay down the proper number on the leaves or scales, and so deter-
mine the phyllotaxy. 1 In a rosette of the leaves of Houseleek
(Fig. 245) and a cone of Pinus Strobus (Fig. 246), the num-
bers which can be seen at one view are appended, and in the
latter the conspicuous secondary spirals are indicated : one to
left with a common difference of 5 ; and two to the right, of
which the most depressed and prominent has the common dif-
ference of 3, the other, nearest the vertical, the common differ-
ence 8. The 14th leaf is superposed to the first, indicating the
Y\ arrangement. The same conclusion is derived from the num-
ber of the higher spirals, the smaller 5 for the numerator, and
this added to 8 for the denominator. The mathematical discus-
sion of these relations, and of the whole subject of phyllotaxy,
leads into interesting fields. But this sketch may suffice for
botanical uses.
246. Relations of Whorls to Spirals. Verticillate and alternate
phyllotaxy, or whorls and spirals, in all complete exemplifica-
tions, are to be considered morphologically as distinct modes,
not to be practically homologized into one. Nevertheless, transi-
tions between the tw r o, and abrupt changes from one to the other
on the same axis are not uncommon, the former especially in
the foliage, the latter in the blossom. If the spiral be assumed
as the fundamental order, it is not difficult to form a clear con-
ception as to how such changes come to pass. A single whorl
1 In applying this method to the determination of the phyllotaxy of a
cone, or any such assemblage of leaves, the student should be warned that,
although the cones of Pines and Firs are all normally on the alternate plan
(while those of Cypresses are on the verticillate), yet in individual cases
(common in Norway Spruce) the cone is plainly made up of pairs of oppo-
site scales which are spirally arranged. See note under 236.
130 PHYLLOTAXY, OR LEAF-ARRANGEMENT.
may most naturally be produced by the non-development of the
intcrnodes between any two, three, or more alternate leaves.
Two proximate distichously alternate leaves would thus form a
pair ; the three leaves belonging to one turn of the spiral in the
tristichous ( J) arrangement would compose a trimcrous whorl ;
the five leaves of the two turns in the pentastichous (|) arrange-
ment, a 5-merous whorl, &c. Verifications of this conception, by
whorls breaking up or reverting to spirals, are occasionally met
with, and the successive overlapping in spiral order of the
members of a trimerous or pentamerous whorl is very common.
The few instances among phaenogamous plants in which the
leaves are opposite and all in the same plane 1 (that is, the suc-
cessive pairs superposed) may be deduced from the distichous
alternate mode becoming opposite without further change, by
the simple suppression of alternate internodes. The frequent
disjunction of the members of the pair in similar and analogous
cases goes to confirm this view. But the characteristic of whorls
ordinaril} 7 is that proximate whorls alternate, that pairs de-
cussate. We cannot homologize this with spiral phyllotaxy ;
for in this lies the fundamental difference between the two plans.
We can explain it only by a reference to Hofmeister's law, which
generally governs leaf-origination as to position, namely, that
succeeding leaves appear directly above the intervals between
the nearest preceding (241, note) : this gives decussation or
alternation of successive pairs or whorls. 2
247. Hypothesis of the origin of both. Instead of regarding
the spiral path on the stem which connects successive alternate
leaves as a purely formal representation, it may be conceived to
be the line along which the members in some original form were
physically connected, in the manner of a leaf-like expansion
1 As in Loranthus Europseus, &c., according to Braun. See 236, note.
2 This renders the verticillate an advantageous arrangement, perhaps no
less so than the distribution which spiral phyllotaxy effects. Both must be
considered to have been determined by and for their respective utilities, and
to have been independent determinations. For " there is no continuity or
principle of connection between spiral arrangements and whorls " (Chauncey
Wright) ; since, although individual whorls are easily reducible to spirals,
each succession is an absolute break of that system.
As whorls of four members often (as especially in calyx, bracts, &c.) may
and sometimes should be viewed as two approximate pairs, so even the spiral
of five members, as in a quincuncial calyx, has been conceived to consist of
two whorls, one of two, the other of three leaves, the second alternating witli
the first as nearly as possible. But this appears far-fetched and of loose
application. It is much clearer as well as simpler to regard the alternate
as the fundamental phyllotaxy, and to deduce individual whorls from spirals,
if need be, rather than to imagine spirals as somehow evolved from whorls.
DISTRIBUTION OF LEAVES OX THE STEM.
131
resembling a spiral stairway. Upon this supposition, the leaves
would be the relics, or rather the advantageous results, of the
segmentation of such a frond-like
expansion, the segments separated
through the development of the
stem in length and firmness, and
modified in the various adaptations
to the conditions of higher vege-
table life ; even as leaves themselves
are modified into tendrils, bud-
scales, petals, or other usefully specialized structures. The type
on this conception would be a frond, consisting of
an elongating axis with a continuous leaf-blade on
one side, and this taking a spirally twisted form.
But the frond of Fucaceous Algae, Hepaticse, and the
like, is two-bladed. While a one-bladed frond, or
with one blade suppressed, might be the original of
alternate-leaved spirals, the two-bladed frond, simi-
larly broken up, would give rise to the opposite or
other varieties of verticillate arrangement. 1
248. Fascicled Leaves need to be mentioned here,
in order that they may be excluded from phyllotaxy.
The}' are simply a cluster or tuft of leaves, belonging
to more than one node, and left in a crowded con-
dition because the internodcs do not lengthen. They
may belong either to the alternate or the verticillate
series. In Barberry and in the Larch (Fig. 247),
they are evidently alternate ; and they may be inferred,
to be so in Pines (Fig. 248), or even may be seen to
be so in the bud-scales which form the sheath sur-
rounding the base of the 2, 3, or 5 foliage-leaves.
In Junipers, the leaves of the fascicles are in the
verticillate order.
1 This is the conception of the late Chauncey Wright. See his elaborate
and most suggestive essay in Mem. Amer. Acad. Arts and Sciences, ser. 2,
ix. 370, mainly reprinted in Philosophical Discussions (posthumous), 296-328,
in which the whole subject of phyllotaxy is acutely discussed, especially
in its relation to questions of origin and developed utilities. His conception
FIG. 247. Piece of a branch of the Larch, with two fascicles of leaves, i. e. two very
short and stout branchlets, bearing scars of former leaves or bud-scales below, and a
dense cluster of leaves of the season at summit. The main axis bears scars from which
the alternate leaves of the developed axis of the preceding year separated.
FIG. 248. Piece of a branch of Pitch Pine, with three leaves in a fascicle or bundle
in the axil of a thin scale (a) which answers to a leaf of the main axis. The bundle is
surrounded at the base by a short shealh, formed of the delicate scales of the axillary
bud, of which the throe leaves are the developed foliage.
132 PHYLLOTAXY, OK LEAF-AIiliANGEMENT.
SECTION II. DISPOSITION OF LEAVES IN THE BUD.
249. Vernation and ^Estivation are terms in general use, under
which the disposition of leaves in the bud is treated. The first
relates to ordinary leaves in this earl}* condition ; the second, to
the parts of a flower-bud ; not, however, as respects insertion,
or position on the axis, which is phyllotaxy (231), but as to
the ways in which they
are coiled, folded, over-
lapped, &c., either perse
O1 '" z ^ r5e - Prcefoliotion
and Prcefloration rre
etymological!}' belter
terms, substituted by
Richard. 1
250. The descriptive
terms which relate to
individual leaves or
parts, whether of foli-
age or blossom, mostly range themselves under the heads of
plications or of enrolling, and are such as the following, the
sectional diagrams of which are copied from the original figures
would make the two plans equally primordial. But the freedom with
which these actually interchange on the same axis greatly favors the lesss
hypothetical view that whorls may be condensed spirals. This assumes
only the well-known fact that internodes may be completely non-developed.
1 Better formed and more expressive terms: but the Linnaean ones are
most in use, and, though fanciful, are not misleading. In English description,
it is as convenient and equally terse to say that the parts are imbricate, val-
vate, &c., " in the bud." Linnseus, in the Philosophia Botanica, described
these dispositions of leaves in the bud under the term Foliatio, not a happy
name, but did not treat of them in the flower-bud. Later, in Termini
Botanici (Amoen. Acad. vi. 1762, reprinted by Giseke in 1781), he intro-
duced the words Vernatio and sEstiratio in their now current botanical sense,
to designate, not the time of leafing and of flowering (spring and summer
condition), but the* disposition of the parts in the leaf-bud and flower-bud
(at least of the petals) as respects foldings, coiling, &c., of single parts, and
modes of overlapping or otherwise of contiguous parts. The terminology
as regards single leaves, Linnaeus fixed nearly as it now remains. That of
leaves or their homologues in connection, and as respects the flower-bud, was
very imperfectly developed until its importance (and much of its termi-
nology) was indicated by Tlobert Brown, in his memoir on Proteaceae, 1809,
in the Prodromus a year later, and in other publications.
Ptyxis (the Greek name) is coming into use as a general term for the
folding, &c., of single parts.
FIG. 249-254. Linmean diagrams <>f sections of leaves In the bud. 249. Condupli?
eate. 250. Plicate or plaited 251. Convojute. 25'J. Revolute. 253. Involute. 254.
Circinate or Circinal
VERNATION, OB PR^EFOLEATION. 133
in the Philosophia Botanica of Linnteus. They were applied
only to foliage, but they are equally applicable to floral parts.
Leaves, and all homologous or similar organs, if not simply
plane, will be either bent or folded or else more or less rolled up
in the bud. The first three of the following terms relate to the
former, the remaining terms to the latter. They are as to the
mode of packing
Plicate or Plaited (Fig. 250), when folded ontlic several ribs,
in the manner of a closed fan, as in Maple and Currant. This
occurs only in certain palmately veined or nerved leaves.
Conduplicate (Fig. 249), when folded lengthwise, or doubled
up flat on the midrib, as in Magnolia ; a very common mode.
The upper face of the leaf is always within.
Redinate or Inflexed, when the upper part is bent on the
lower, or the blade on the petiole, as in the Tulip-tree (the blade
of which is also conduplicatc) .
Convolute (Fig. 251), when rolled up from one margin, i.e.
one margin within the coil, the other without, as in Apricot
and Cherry.
Involute (Fig. 253), botli margins rolled toward the midrib
on the upper face, , as the leaves of Water Lily, Violet, &c. ;
also the petals of Steironema and Treinandra.
Revolute (Fig. 252), similarly rolled backward from both
margins, as the leaves of Azalea and Rosemary.
Circinal or Oircinate (Fig. 254), when coiled from the apex
downward, as the leaves of Drosera and the fronds of all the
true Ferns.
Corrugate or Crumpled, as the petals of a Poppy, applies
to the irregular crumpling of the otherwise plane corolla-loaves.
This is a consequence of rapid growth in length and breadth
in a confined space.
251. The Ptyxis (or folding, &c.) of an individual leaf, of
which the foregoing modifications are the principal, should be
distinguished from the arrangement in the bud of the leaves of
a circle or spiral in respect to each other. The interest of the
latter centres in the flower-bud, i.e. in aestivation. To this the
following exposition is devoted, although sometimes applicable
to leaf-buds -also. 1
252. The disposition of parts in aestivation, in respect to
each other, is the result partly of their relative insertion, that is
1 In the succeeding paragraphs, it becomes necessary to presuppose so
much knowledge of the flower as is implied in the free use of such terms as
calyx and corolla, sepals or calyx-leaves, and petals or corolla-leaves. See,
if need he, Chapter VI. Sect. I.
134 PHYLLOTAXY, OR LEAF-AHRANGEMENT.
their phyllotaxy, and partly of the way in which they comport
when their margins meet in growth. Those leaves which are
within, or of higher insertion on the axis, will almost necessarily
be enclosed or overlapped : those which are members strictly
of the same whorl or cycle may fail to come into contact, or
may meet without overlapping at the contiguous margins or
apex ; yet they may be overlapped, since they may have grown
unequally or some a little earlier than their fellows. Conse-
quently, no perfectly clear line can be drawn in the flower between
cycles and spirals except by their mode of succession. More-
over, {estivation strictly so called should be concerned only
with the disposition among themselves of the several members
of one whorl, or of one complete spiral. So the alternation of
contiguous whorls, as of the three inner with the three outer
flower-leaves of a Lily or a Tulip (the alternative {estivation of
DeCandolle), is a matter of phyllotaxy, not of aestivation. The
latter is properly concerned only with the relations of each three
loaves to each other. 1
253. The proper aestivations ma}' be classified into those in
which the parts do not overlap, and those in which they do. Of
the first, there are two kinds, the open (cest.
aperta) and the valvate, both characterized and
nanu'd by Brown. 2 Of the second, there is
one leading kind, the imbricate (adopted by
Brown from Linnaeus), with subordinate modi-
fications. 3 Accordingly, the aestivation is
said to be
2 ~> 1. Open or Indeterminate (cesf. aperta)^
when the parts do not coma into contact in the bud, so as to
1 The same applies to the two sets of sepals and of petals in Barberry, in
Menispermum, and of the petals in Poppy, &e. (050).
' 2 Linnaeus, indeed, has, " sEstiratio vaJvata, si petala se expansura instar
gluma? graminis ponuntur," the name, but not the thing: the glumes of
grasses are not valvate in the botanical sense. So the term as to its proper
use maybe said to originate with "R. Brown.
3 For a brief discussion of "^Estivation and its Terminology," see Amer.
Jour. Sci. ser. 3, x. 339, 1875.
As to names, it is perhaps more correct to say of the (t'atii'ation that it is
inihricat/ve, convotutire, valvular, &c. (), pedicels (p'), bracts (b). and
bractlete (V).
144 ANTHOTAXV, OK INFLORESCENCE.
266. Position of Flowers or Clusters. Flower-buds accord
with leaf-buds in origin, position, and structure, to this extent at
least, that the parts of both are leaves or homologues of leaves,
crowded in whorls or spirals upon a short portion of stem or
axis; and as leaf-buds are either terminal or axillary (15, 75)
so also are flower-buds ; as a leaf-bud may give rise to a simple
or a compound growth, i. e. may branch again and again, or not
branch at all, so flower-bearing branches, or the flower-bearing
extremit}' of a stem or branch, may bear a single flower, or a
more or less compound cluster. Thus, in Fig. 276, an axillary
peduncle, or naked branch, bears at apex a solitary flower ; in
Fig. 277, a peduncle bears a loose cluster of flowers, each of
which springs from the axil of a small bract ; in Fig. 285,
a terminal peduncle bears at summit a dense flower-cluster.
Flowers are either solitary or in clusters. When solitary, they
are naturally without bracts, being subtended instead (as in Fig.
276) by ordinary foliage.
267. The elevation either of a solitary flower or a cluster on a
peduncle, or of individual flowers of a cluster on pedicels, is only
incidental. The flowers may be stalkless, i. e. sssil(>.
268. The Kinds of Inflorescence which have received distinctive
names are various, but are all reducible to two types, which,
generally well marked, may sometimes pass into each other, and
which are not rarely combined in the same compound inflores-
cence. 1 The two types differ in basis as do axillary from ter-
minal buds ; in the one the flowers are axillary or lateral, in the
other terminal in respect to the axis from which each flower or
its pedicel arises. But inasmuch as ever}' flower, whatever its
position, is terminal to its own stalk or axis, it is better to dis-
tinguish the two types in other terms, and to name them the
269. Indefinite and Definite, or, in equivalent and similar terms,
the Indeterminate and Determinate. 2 Each ma} r be either simple
or compound. It is from the simple that the definitions arc to
be drawn. In the former type, the rhachis or main axis of the
inflorescence is not terminated by a flower, but lateral axes, or
pedicels, are. In the latter, both the main or primary and the
lateral or secondary axes or stalks are so terminated. An inde-
terminate flower-cluster may go on to develop internode after
1 Inflorescence, as has been well insisted on by Guillaml (in Bull. Soc.
Bot. France, iv. 29), is a mode, not a thing. The things sometimes but in-
appropriately so called are flower-clusters, for which, if a general technical
name is needed, that ofAnthemia, in English Anthemy, suggested by Guillaud,
is as good as any.
? - Also named by Eichler (Bliithendiagramme, 33, following Guillaud, 1. c.)
the Cyntose and the Botryose type.
THE TWO TYPES.
145
internode of axis, and one or more leaves (bracts) at each node,
and then a flower in the axil of each bract, until its strength or
capability is exhausted. Or it may stop short with very few
flowers ; but the uppermost and youngest one will not realty ter-
minate the rhachis (i. c. come from a terminal bud) , though it
may appear to do so. (Fig. 272, 277-279, &c.) The lower
flower-buds are evidently the oldest, and accordingly the first to
expand ; and the expansion will proceed regularly from below
upward : wherefore this type of inflorescence has been called the
Ascendiny or Acropetal ; likewise the Centripetal, because, when
the flowers are brought to the same level or near it (as in Fig.
279, 280) by a lengthening of the lower pedicels, with or with-
out relative shortening of the rhachis, the evolution
jj\ is seen to proceed from circumference to centre.
\| ($ &fr There is thus no lack of names ; but, inasmuch as
A J the following type is commonly referred to under the
Mfe general name of Cymose, to this has recently been
given the counterpart name of Botryose. (271.)
F
281 282 283 284
270. A determinate flower-cluster (as seen in its gradual
development which is not rarely presented) has the last internode
FIG. 278-280. Diagrams of indefinite, indeterminate, centripetal, or botryose in-
florescence: 278, Raceme; 279, Corymb; 280. Umbel.
FIG. 281-284 Diagrams of definite, determinate, centrifugal, or cymose inflores-
cence : 281, a false or descending raceme ; 282, a solitary terminal flower ; 283, same with
two lateral flowers developing, forming a 3-flowered cyme; 284, same with lateral
pe'luncles 3-fl'>wered, or a pair of 3-flowere 1 cymes, beside the central or primary
terminal flower.
146 ANTHOTAXY, OK INFLORESCENCE.
of its axis terminated by a flower (Fig. 281-284), which answers
to a terminal bud. If more flowers appear, so as to compose a
cluster, the}' spring from the axils, preferably from the highest
axils, and are later. The order of evolution is shown in the figures
by the size of the flower-buds or degree of expansion of the
blossoms. Fig. 281 best shows why a determinate or definite in-
florescence is sometimes said to be Descending ; Fig. 283 shows
why it is called Centrifugal, the central flower first expanding ;
Fig. 284 exhibits the lateral or circumferential partial clusters
later than the central blossom, and their lateral flowers later
than their central.
271. Varieties of Indeterminate or Botryose Inflorescence. The
names of most of these have been fixed from the time of Linnaeus,
but defined without reference to the order of evolution of the
flowers. They are the Raceme, Corymb, and Umbel, with flowers
raised on pedicels ; the Spike and Head, with sessile flowers ;
also some modifications of these, notably the Ament and the
Spadix. The raceme may be taken as the type. Botrys is
equivalent to racemus, &c. ; and, as the type includes diversity
of forms to which the name racemose would seem inapplicable,
the term botryose (botrytischen of Eichler) is best chosen as the
general name of it, and is a good counterpart to cymose for the
other type.
272. A Raceme (illustrated in Fig. 272, 277, and by diagram
in Fig. 278) is a simple flower-cluster, in which the flowers,
on their own lateral or axillary pedicels and of somewhat equal
length, are arranged along a relatively more or less elongated
rhachis or axis of inflorescence. The common Barberry, Cur-
rant, Choke-Cheny and Black Cherry, and Lily of the Valley
are familiar examples.
273. A Corymb (Fig. 275, 279) is a shorter and broader
botiyose cluster, which differs from a raceme only in the relatively
shorter rhachis and longer lower pedicels ; the cluster thus be-
coming flat-topped or convex. The centripetal character is thus
made apparent. The greater number of the corymbs of Linnaeus
and succeeding botanists are cymes, the central flower first ex-
panding. And the term corymbose or corymb-like is still much
used in descriptive botany for a ramification which is mainly of
the cj'mose tj*pe, and where in strictness the term cymose should
be employed.
274. An Umbel (Fig. 280), as in Asclepias, &c., differs from
a corymb only in the extreme abbreviation of the rhachis or axis
of inflorescence, and the general equality of the pedicels which
thus all appear to originate from the apex of the peduncle, and
THE BOTRYOSE TYPE.
147
so resemble the ra}*s of an umbrella; whence the name^and
whence also the pedicels or partial peduncles of an umbel are
termed its Rays. The bracts, brought by the non-development
of internodes into a depressed spiral or apparent (or sometimes
real) whorl, become an involucre. (264.) An umbel or any
similar cluster when sessile (without a common peduncle) , and
the parts crowded, is sometimes called a Fascicle (or the pedicels
said to be fascicled) ; but this term has been differently defined.
(280.) It is better not to use it for any special kind of inflores-
cence, but simpl}* in the sense of a bundle of whatever sort.
This will accord with the sense in which it is applied to an
aggregation of leaves. (248.)
275. A Head or Capitulum (Fig. 285) is a globular cluster of
sessile flowers, like those of Red Clover, Button-bush, and
Plane-tree. The pedicels need not be absolutely wanting, but
only very short. An umbel
with pedicels much abbrevi-
ated thus passes into a head,
as in Eryngium, &c. And
a head with rhachis elongated
passes into a spike. The
short rhachis of a head very
commonly takes the name of
receptacle. (205.) The whole
may be subtended by con-
spicuous bracts forming an
involucre (264) as in Fig.
286, or may be destitute of
any, as in Fig. 285. On ac-
count of the compactness
and mutual pressure under
growth, the bracts among
the flowers of such heads (normally one subtending each blos-
som) are apt to be rudimentary, reduced to little scales, or
abortive, or completely wanting. In the latter case, the recep-
tacle is said to be naked (nude), i.e. naked of bracts: when
they arc present, it is pakate or chaffy. A peculiar sort of head,
not undeserving a special name (though this is not necessary
in descriptive botany) , is the
ANTIIODIUM, the so-called Compound Flower of the earlier
botanists, which gives the name to the vast order of Composite.
FIG. 285. Cephalanthus ocndcntalis, the Button-bush; a pair of leaves, and a
terminal peduncle bearing a dense head (capitulum) of flowers.
148 ANTHOTAXY, OR INFLORESCENCE.
The,, name means "resembling a flower." Although it has all
the characters of a true head, the resemblance to a flower is
remarkably striking, the involucre imitating a calyx, and the
strap-shaped (ligulate) corollas of the several flowers imitating
the petals of a single blossom. In some (such as Dandelion
and the Cichory, Fig. 286) , all
the flowers of the head bear
these petal-like corollas ; in
more (such as Aster, Sun-
flower, and Coreopsis, Fig.
287) , only an outer circle of
flowers does so ; the remain-
der, smaller and filling the
centre (or disk), may by the
casual observer be taken for
stamens and pistils, and further the deception. The rhachis
or receptacle of a head of this kind is commonly depressed,
bearing the flowers on what then becomes the upper surface,
which adds to the imitation. 1
SYCONIUM. This name, given to the Fig-fruit, should be here
referred to, as it is a sort of inflorescence, of the general nature
of a head, but with receptacle external and flowers enclosed
1 The receptacle of an Anlhodhtm has been termed Clinanthium or Phor-
anthium ; and its involucre, a Pfripkvraidkiam or Peridinium. The head has
likewise been named a Cephalanfhium.
FIG. 286. Flowering branch of Cichory, with two heads of ligulate flowers, of
natural size.
FIG. 287. Vertical section of a head of flowers of a Coreopsis.
THE BOTRYOSE TYPE.
149
within. See Fig. 657-659 (683), where its morphology is ex-
plained and illustrated. Viewed as an inflorescence, it has also
been named a HYPANTHODIUM.
276. A Spike is a cluster of sessile (or apparently or nearly
sessile) lateral flowers on an elongated axis. It may be de-
fined by comparison, as a head with the rhachis lengthened
(indeed a young head often becomes a spike when older), and
equally as a raceme with the pedicels
all much shortened or wanting. A
common Mullein and a Plantain
(Plantago, Fig. 290) are familiar ex-
amples. Two modifications of the
spike (or sometimes of the head) gen-
erally bear distinct names, although
not distinguishable by exact and con-
stant characters, viz. :
SPADIX, a spike or head with a
fleshy or thickened rhachis. The
term is almost restricted to the Arum
family and Palms, and to cases in
which the inflorescence is accompanied
by the peculiar bract or bracts called
a spathe (Fig. 269-271). But the
two do not always go together : in
Acorns and Orontium there is properly
no spathe to the spadix ; while in the
Iris family the bracts are said to form
a spathe, and there is no spadix. In Palms, the principal reason
for naming the inflorescence a spadix is its inclusion in a
spathe before anthesis.
FIG. 288. A slice of Fig. 287. more enlarged, with one tubular perfect flower (a) left
standing on the receptacle, and subtended by its bract or chaff (ft); also one ligulate
and neutral ray-flower and part of another (c, c): in d, d, the bracts or leaves of the
involucre are seen in section.
FIG. 289. Catkin of White Fireh. 290. Young spike of Plantago major.
150
ANTHOTAXY, OR INFLORESCENCE.
AMENT or CATKIN. This is merely that kind of spike with
scaly bracts borne by the Birch (Fig. 289), Poplar, Willow, and,
as to one sort of flowers, by the Oak, Walnut, and Hickory,
which are accordingly called amentaceous trees. Catkins usually
fall off in one piece, after flowering or fruiting. All true catkins
are unisexual.
277. Any of these forms of simple inflorescence ma}' be com-
pounded. Racemes may themselves be disposed in racemes, spikes
in spikes (as in Triticum) , heads
be aggregated in heads, umbels in
umbels, corymbs ma} 7 be corym-
bosely compound, &c. ; forming
compound racemes, spikes, umbels,
and the like, the terminology of
which is easy. The most usual
case of trul}' homomorphous com-
pounding is that of umbels ; the
inflorescence of much the larger part of Umbclliferae being in
compound umbels, as in Fig. 290*. There is then the general
umbel, the ra} T s of which become peduncles to
the partial umbels, and the rays of the latter
are pedicels. Umbella and Umbellulce desig-
nate in Latin terminology the general and its
partial umbels. Umbellets (coined by the late
Dr. Darlington) may well replace the latter
as the English diminutive. But umbels are
sometimes racemosely arranged, as in Aralia
spinosa, heads may be arranged in spikes,
and so on.
278. A Panicle, of the simple and normal
sort (as illustrated in Fig. 291), is produced
when a raceme becomes irregularly compound
by some (usually the lower) of its pedicels
developing into peduncles canying several
flowers, or more than one, or branching again
and again in the same order. But in com-
pound clusters generally the secondary and
tertiary ramifications are apt to differ in t}*pe
as well as in particular mode, giving rise to
heteromorphous or mixed inflorescence. (288.)
As Linnaeus defined the term, and as it has generally been em-
ployed in botanical descriptions, the panicle is a general term
FIG. 290<. Compound umbel of Caraway. 291. A simple panicle.
THE CYMOSE TYPE.
151
for any loose and diversely branched cluster, with pedicellate
flowers. It is therefore difficult to restrict it in practice to the
indeterminate type.
279. Varieties of Determinate or Cymose Inflorescence. The
plan of this type has been sufficiently explained. (270.) Its
simplest condition is that of a solitary terminal flower, peduncu-
late or pedicellate (as in Fig. 282), or sessile. The production
of more flowers necessitates new axes from beneath, from the
axils of adjacent leaves or bracts. These, being later, render
the evolution centrifugal. The simplest flower-cluster (unless
we call the solitary flower of Fig. 282 a one-flowered cluster) is
that of Fig. 283, where a secondary floral axis or peduncle has
developed from the axil of each leaf of the uppermost pair, or
where with alternate leaves there is a single uppermost leaf, and
then only one such peduncle, and thus is produced a three- (or
two-) flowered cymose cluster. The flower of the primary axis
is marked by its bractless peduncle (therefore a pedicel) ; the
lateral and secondary peduncles are known (commonly or nor-
mally) by their bracts or bract ; the portion below the bracts
is proper peduncle ; that above, of single internode, pedicel.
Bracts, like other leaves, have potential buds in their axils ; these
in an inflorescence give the third order of ramification, each
branch tipped with its flower ; and so on.
280. The Cyme is the general name of this kind of flower-cluster
in its various forms. One of these very simple c}'mes, by itself or
as a part of a larger cyme, may be called a Cymule. The regular
cj'mc usually accompanies opposite or other grades of verticillate
leaves, but is not rare in the alternate arrangement. It is
readiest understood in an opposite-leaved plant with regular
opposite ramification, as in an Arenaria, Fig. 292. By its con-
stitution, a cyme proceeds from simple to compound. It mat-
FIG. 292. Dichotoiuous or Mparous cyme (cyme bipare of Bravais, Dichasium of
Eichler) of Arenaria Michauxii.
152 ANTHOTAXY, OR INFLORESCENCE.
ters little whether its development is progressive, the flowers of
the ultimate ramifications expanding after the earlier have matured
fruit, and with subtending bracts conspicuous or foliaceous ; or
whether, as in Elder and Hydrangea (Fig. 293, and in Fig.
273), the bracts are minute and caducous or abortive, and the
ramification complete with all the flower-buds well formed before
the oldest expand, so that the whole is in blossom almost at the
same time. But a cyme may be properly said to be compound
when the primary axis in it is a peduncle instead of a pedicel,
M
and supports a cluster (cyme or cymule) instead of a solitary
central flower at the main divisions. 1 One form of the regular
cj r me, on account of its compactness, is named the
GLOMERULE. This is merely a cymose inflorescence, of any
sort, which is condensed into the form of a head, or approach-
ing it. Of this kind is the so-called head of Cornus florida, and
of the herbaceous C. Canadensis (Fig. 294), which shows the
1 The dieliotomous or two-branched cyme is the commonest, but is some-
times marked by suppression of internodes ; as, for example, where the
branches are apparently in fours, in an umbelliform way ; but these are two
sets of two, with the internode between the pairs extremely short ; or where,
as in Elder, the branches or rays are five, in this case consisting of the same
two pairs and a central one, which is a many-flowered continuation of the
primary axis. Or 5-rayed cymes, &c., may be founded upon alternate leaves
with shortened internodes, the rays or peduncles axillary to them thus
brought into an apparent whorl.
Bravais distinguished cymes as multiparons, with three or more lateral
axes ; biparous, with two ; and uniparons, with only one (cyme multipare,
bipare, unipare). To these Eichler gives the substantive names, severally,
of Pleiochasium, Dichasium, and Monochasium. Only the latter needs illustra-
tion ; the others being as it were compounds of this.
FIG. 293. Compound cyme of Hydrangea; with some neutral and enlarged mar-
ginal flowers.
THE CYMOSE TYPE.
153
composition best, on close examination. A condensed but less
capitate cyme, or cluster of cymes, was called by Keeper and
DeCandolle a FASCICLE ; and this terminology has been much
adopted. It is properly enough
said to be a fascicle, which, as
used by Linnaeus and others,
means a bundle, or close collection
of parts, whether leaves, pedun-
cles, or flowers ; but a fascicle is
not necessarily a cyme (274) , nor is
there need of a special substantive
name for a compact cyme, which
may either be simply so called or
it may pass into the glomerule.
281. Botryoidal forms of Cymose
Type, or False Racemes, &c. The
regular cyme seldom continues
with all its ramifications. In
Fig. 292, after the second forking,
one of the two lateral peduncles
mostly fails to appear, and in some
parts one of the bracts also ; and ultimately the lateral peduncle
present is bractless, like the central, therefore equally incapable
of further ramification, being reduced to a pedicel of a single inter-
node. This suppression some-
times begins at the first fork-
ing or at the very base ; and,
when followed throughout, it
reduces a biparous or dichoto-
mous C3'me to one half, and,
converts this half (when the
axis straightens) into the sem-
blance of a raceme if the
flowers are pedicelled, or of a
spike when the} r are sessile.
Fig. 296 is a diagram of such an inflorescence as that of Fig. 292,
with one lateral branch uniformly suppressed at each division, the
wanting members indicated by short dotted lines. Cases exem-
plifying this occur in portions of the inflorescence of some of our
FIG. 294. Plant of Cornus Canadensis: flowering stem bearing a cluster of leaves
above, then continued into a peduncle, and terminated by a glomerule of very small
flowers; this subtended by a colored and corolla-like involucre of four bracts. 295 One
of the flowers taken from the glomerule. enlarged.
FIG. 296. Uniparrms cyme or sympodial false raceme, with opposite leaves or bracts.
FIG. 297. Form of the same, with alternate leaves or bracts.
154 ANTHOTAXY, OR INFLORESCENCE.
smaller ttyperictims, and notably in H. Sarothra, in which the
leaves are all reduced to bracts. It is not always easy to show
why this is not a true raceme. But the other bract of the pair, upon
that supposition, is unaccountably empty : the successive angular
divergence of each joint of the axis of inflorescence in the younger
part, which commonly runs into a coil, finds explanation in the view
that each portion is the lateral branch from the axil of the subtend-
ing leaf: and occasionally the other axil produces a similar one,
thus revealing the cymose character. When the bract from the
axil of which the missing branch should come disappears also, as
sometimes it does, and uniformly on the same side, a state of
things like that of the upper part of Fig. 29 7 occurs. The same
figure may serve for the arrangement corresponding to that of
Fig. 296, only with alternate leaves. But then, close as the imi-
tation of a raceme here is, the position of each flower in respect
to the bract supplies a criterion. While in a true raceme the
flower stands in the axil of its bract, here it stands on the oppo-
site side of the axis, or at least is quite away from the axil.
282. Sympodial forms. The explanation is that the axis of
inflorescence in such cases, continuous as it appears to be, is
not a simple one, is not a monopode, but a sijmpode (110, 116,
notes), i.e. consists of a series of seemingly superposed inter-
nodes which belong to successive generations of axes : each axis
bears a pair of leaves (Fig. 296) or a single leaf (Fig. 297),
is continued beyond into a peduncle (or pedicel in these
instances), and is terminated by a flower. From the axil
promptly springs a new axis or branch, vigorous enough soon
to throw the adjacent pedicel and flower to one side : this
bears its leaf or pair of leaves, and is terminated like its prede-
cessor with a flower ; and so on indefinitely. The fact that the
alternate leaves or bracts are thrown more or less strictly to one
side and the flowers to the other, in Fig. 297, shows that these
leaves do not belong to one and the same axis ; for alternate
leaves are never one-ranked or disposed preponderatingly along
one side of an axis, as in this diagram, and as is seen in the
inflorescence of a Houseleek, &c.
283. A further difficulty in the morphology of clusters of this
class comes from the early abortion or complete suppression of
bracts. This is not unknown in botryose inflorescence, occurring
in the racemes of almost all CrucifenE : it is very common in the
cymose of all varieties, and especially in the uniparous ones in
question, which characterize or abound in Borraginaceae, Hydro-
plryllaceae, and other natural orders. In some genera or species,
the bracts are present, or at least the lower ones ; in others,
THE CYMOSE TYPE. 155
absent ; in some, either occasional!}' present or wanting in the
same species or individual. It is only by analogy, therefore,
and by a comparison of allied plants, that the nature of some of
these flower-clusters can be made out. With the botanists of a
preceding generation, tlxese one-sided clusters were all described
as racemes or spikes. Botanists still find it convenient to con-
tinue the use of these names for them in botanical descriptions,
adding, however, as occasion requires, the qualification that they
are false racemes or spikes, or cymose racemes, and the like ; or
else, bj- reversing the phrase, with stricter correctness they call
them racemiform or spiciform cymes, &c.
284. Commonly these false racemes or spikes (or botiyoidal
C}'mes, if we so name them) are circinate or inrolled from the
apex when young, in the manner of a crosier, straightening as
they come into blossom or fruiting. Likening them to a scorpion
when coiled, the earlier botanists designated this as scorpioid.
As the coil is a helix, it has also been named helicoid. 1 The
flowers are then thrown, more or less strictly, to the outer side
of the coiled rhachis, where there is room for them ; and so these
false racemes or spikes are secund or unilateral. The particular
anthotaxy and phyllotaxy of the various s^ympodial and botryoi-
dal forms of cymose inflorescence become rather difficult ; and
the sorts which have been elaborate!}* classified into species
(and have no little morphological interest) are connected by
such transitions, and are based on such nice or sometimes theo-
retical particulars, that the terminology based on them is seldom
conveniently applicable to descriptive botany, at least as to sub-
stantive names.
285. One of the latest and simplest classifications of C3*mes is
that of Eichler in his Bluthendiagramme. 2
1 Scorpioid and Helicoid have been carefully distinguished by later
morphologists, on account of some difference in the mode of evolution and
arrangement of the flowers along one side of the rhachis, by which they
become two-ranked in scorpioid, one-ranked in helicoid. But practically
the two kinds of clusters are not always readily discriminated ; and in gen-
eral terminology a single name, with subordinate qualifying terms, is suffi-
cient. Scorpioid is the older and commoner one, therefore the most proper
to be used in the generic sense.
2 CYMOSE TYPE (classified without reference to bracts, which are so often
wanting) ; divided into
o. Lateral axes three or more: PLEIOCHASIUM, the multiparous cyme ot
Bravais.
)8. Lateral axes two: DICHASIUM, the biparous cyme of Bravais.
7. Lateral axis one: MONOCHASIUM, the uniparous cyme of Bravais.
The latter, or the corresponding divisions of the preceding sorts, may be
divided as follows :
166
ANTHOTAXY, OR INFLORESCENCE.
286. Sundry complications and obscurities are occasionally
encountered in anthotaxy or phyllotaxy, which cannot here be
* Lateral axes transverse to the relatively main axis.
1. Lateral axes in successive generations always falling on the same
side of the relatively main axis : SCHRAUBEL [screwlike] or BOSTRYX
[ringlet or curl], the uniparoits helicoid cyme of Bravais.
2. Lateral axes falling alternately on opposite sides of the relatively main
axis: WICKEL or CINCINNUS [a curl], the uniparous scorpioid cyme of
Bravais.
* * Lateral axes medial [in the same plane] relative to the main axis.
3. Lateral axes in successive generations always on the back side of the
axis from which it springs: FACHEL, RHIPIDIUM [fan].
4. Lateral axes in successive generations always on the upper side of the
axis from which it springs : SICHEL, DREPANIUM [sickle].
The subjoined simple diagrams from Eichler (Fig. 298-303) illustrate
these forms. The ramification is given without the bracts, which theoreti-
cally or actually subtend the axes of each generation. The student may add
them, and so more readily apprehend the characters.
Eichler recognizes the forms with median (antero-posterior) position of
axes in Monocotyledons only.
It is natural to distichous phyl-
lotaxy, and it accords with
the general rule that, in mono-
cotyledonous plants, the first
leaf of the branch, or the com-
monly solitary bractlet of the
|6 peduncle, stands over against
and facing the bract or leaf
from the axil of which said
branch or peduncle springs, i. e. is posterior and next the parent axis, as
shown in the diagram, Fig. 304, 305.
301 300 304
a a'
a b' a' b
W
FIG. 298. Diagram of the Cincinnus. 299. Diagram of the Bostryx. The flower-
axes numbered in succession.
FIG. 300. Diagram of the Rhipulium. 302. Ground plan, indicating the order of
evolution of the flowers.
FIG. 301. Diagram of the Drepanium. 303. Ground plan, the flowers evolved in
succession, from left to right.
FIG. 304. Diagram showing the position of bractlet or first leaf on a branch in
Monocotyledons: a is the primary, a f the secondary axis*; b is brack and I' bractlet.
THE CYMOSE TYPE. 157
explained, except through full details : such as flowers standing
by the side of a leaf, or a small leaf b} r the side of a larger one,
The transverse or oblique position of secondary axes or peduncles, as in
Eichler's first two species, brings the flowers of the false raceme or spike
out of line of the sympodial axis and bracts, neither in the axils, as in true
racemes, nor opposite them, as in the Rhipidium and Drepanium, but on one
side of this plane or the other. This is most common in Dicotyledons (in
Drosera, Sedum, Sempervivum, and Hyoscyamus, in Borraginacese and
Hydrophyllaceae, &c.), and is not rare in Monocotyledons, especially with
tristichous phyllotaxy, as in Tradescantia. In the Bostryx, Fig. 299, the
bractlet is anterior or falls on the same side as the bract, or, in other terms,
the successive bracts are all on one side, the inner side, of the helix ; and the
Drepanium (Fig. 301) is like it: this is the helicoid cyme of Bravais, &c.,
and its flowers are commonly one-ranked. In the Cincinnus or true scorpioid
cyme (Fig. 298), and equally in the Rhipidiurn (Fig. 300), the bracts fall
alternately on opposite or different sides of the sympodial rhachis, because
the single bract (b f ) of each successive secondary axis (a f ) stands next the
axis (a) and over against the bract (b) of the generation preceding. The
flowers in these generally fall into two parallel ranks (conspicuously so
when crowded) on the upper side of the rhachis, on which, in the cincinnus
or true scorpioid cluster, they are usually sessile or nearly so (or spicate),
as is well seen in Heliotropes, and in very many Borraginaceous and Hydro-
phyllaceous species, in Houseleek, Tradescantia erecta, &c. This comes
through antidromy, that is, the phyllotaxy of
each successive axis of the sympode (with its
one bract, or by suppression without it) changes
direction, from right to left and from left to
right alternately. Fig. 301 is a plan of this
two-ranked unilateral arrangement. When not
too crowded, both Cincinnus and Rhipidium
are apt to have a zigzag rhachis.
These two last-mentioned kinds are so gen-
erally alike in character, as are equally the
Bostryx and the Drepanium, that the four spe-
cies may as well be reduced to two. As these
severally include the scorpioid and the helicoid
uniparous cymes of modern anthotaxy, these
terms may be retained to designate them. Or,
if other terms in use be preferred to scorpioid
ami helicoid, the form with two-ranked flowers
may be denominated Cincinnal, that with single-
ranked Bostrychofdal. But in neither type is the rhachis always coiled
up, although commonly more or less so in the undeveloped state.
While these forms generally imitate racemes or spikes, it will be noted
that Fig. 300 specially imitates a corymb in form and in seeming acropetal
or centripetal evolution. And when, as in this figure, the bracts are all
absent, no obvious external difference remains.
FIG. 305. Ground plan (from Eichler's Bluthendiagramme, i. 38) of the scorpioid
inflorescence of Tradescantia erecta, between bract (B) below and axis (a) above:
I., II., III., &c., the successive flowers: v l is the bractlet of the first and bract of the
second flower, and so the others in succession up to v 5 and a small undeveloped one
beyond. The figure 1 affixed to each flower indicates the first floral leaf.
158 ANTHOTAXY, OR INFLORESCENCE.
or a pedicel or peduncle above and out of connection with the
leaf which should subtend and accompairy it. 1
287. Mixed Inflorescence is not uncommon. This name is
given to clusters or ramifications in which the two types are con-
joined. Being heteromorphous, they are almost necessarily com-
pound, the two types belonging to different orders of ramification.
But under it may be included cases of comparatively simple
inflorescence, at least in the beginning, some of which nearly
fuse the two types into one. In the Teasel (Dipsacus) , an appar-
ently simple head or short spike comes first into flower at the
middle, from which the flowering proceeds regularly to the base.
Had it begun at the top, it would answer to Fig. 281, which,
blossoming from above downward by simple uniflorous lateral
axes along a monopodial primary axis, is a simple racemiform
C3*mc, while it may also be called a reversed or determinate raceme.
Something of this sort may be seen in certain species of Cam-
panula, with virgate inflorescence, the terminal blossom earliest,
the others following irregularly, or parti}' downward and partly
upward. In C. rapunculoides, when rather depauperate and
the inflorescence simple, the evolution is that of a true raceme,
except that a flower at length terminates the axis and develops
earlier than the upper half of the raceme. In Liatris spicata
and its near relatives, the heads, on the virgate general axis,
come into flower in an almost regular descending order, or are
reversely s/ncate. If in Fig. 281 the lower pedicels were prolonged
to the level of the upper, a simple eorymbiform cyme would be
seen, with simple centrifugal evolution, that is, regularly from
the centre to the margin ; this is the counterpart of the rhipidium
or fan-shaped c}'me, of Fig. 300, in which the evolution of the
blossoms is as regularly centripetal. The explanation of the
paradox is not far to seek.
1 The position of a pedicel at the side of a bract in false racemes is ex-
plained in the foregoing note. It may occur in true racemose inflorescence
by the reduction of sessile secondary racemes down to an umbel of two
flowers, transverse to the bract (as in many species of Desmodium), and
thus seemingly lateral to it, or to a single flower on the right or left of it.
The coalescence of a pedicel to the axis for a considerable height above the
subtending bract in a simple inflorescence, or above the last leaf in a
sympodial one (concaulescence of Schimper), is common. So likewise bracts
or leaves may be for a good distance adnate to sympodial shoots, whether
peduncles or leafy flowerless branches. This (named recaulrscence by
Schimper) is of most frequent occurrence in Solanaceae (in Datura, Atropa,
most species of Solan um, &c.), and is the explanation of their so-called
geminate leaves, where a large leaf (really belonging lower down) has a
small leaf by the side of it. See Wydler in Bot. Zeit. ii. 689, &c., Sendtner
in Fl. Bras. x. 183, and Eichler, Bluthend. i. 199.
THE CYMOSE TYPE. 159
288. Compound mixed inflorescence is very various and com-
mon ; but the combinations have hardly called for special terms,
being usually disposed of by a separate mention of the general
and of the partial anthotaxy, or that of the main axis of inflo-
rescence and that of its ensuing ramification. 1 In Composite,
for instance, the flowers are always in true heads, of centripetal
evolution. The heads terminate main stems as well as lateral
branches, so that they are centrifugally or C3'mosely disposed.
The reverse occurs in all Labiatse and most Scrophulariaceae,
where the flowers, when clustered, are in cymes, but these cymes
are from axils, and develop in centripetal order. It is this
arrangement which mainly characterizes the
THYRSUS. A compound inflorescence of more or less elongated
shape, with the primary ramification centripetal or botryose, the
secondary or the ultimate centrifugal or cymose. To the defini-
tion is generally added, that the middle primary branches arc
longer than the upper and lower, rendering the whole cluster
narrower at top and bottom, and sometimes that it is compact ;
but these particulars belong only to typical examples, such as
the inflorescence of Lilac and Horsechestnut. In the former, the
thyrsus is usually compound. A loose thyrsus is a
MIXED PANICLE. It is seldom that a repeatedly branching
inflorescence of the paniculate mode is of one t} r pe in all its
successive ramifications. Either the primarily centripetal will
become centrifugal in the ultimate divisions, or the primarily
centrifugal will by suppression soon run into false botryose
forms, into apparent racemose or spicate subdivisions. So that
the name Panicle in terminology is generally applied to all such
mixed compound inflorescence, as well as to the homogeneously
botryose. (278.)
VERTICILLASTER is a name given to a pair of opposite and
sessile or somewhat sessile cymes of a thyrsus or thyrsiform
inflorescence, which, when full, seem to make a kind of verticil or
whorl around the stem, as in very many Labiatae. The name
was originally given to each one of the pair of cymes ; but it is
better and more commonly used to denote the whole glomerule
or false whorl produced by the seeming confluence of the two
clusters into one which surrounds the stem.
1 Guillaud (in his memoir on Inflorescence, published in Bull. Soc. Bot.
France, iv.) proposes to designate as Ci/mo-Botryes the mixed inflorescence
composed of cymes developed in botryose order, i. e. the thyrsus ; and
Botry- Cymes, the reverse case of racemes, &c., cymosely aggregated. For
the former, the old name thyrsus serves appropriately and well.
160 ANTHOTAXY, OR INFLORESCENCE.
289. The Relations of Bract, Bractlet, and Flower should here
be noticed, although the subject in part belongs rather to the
section on Floral Jyymmetiy. (315.)
290. Anterior and Posterior, otherwise called Inferior and
Superior, and therefore Lower and Upper, 1 are primary relations
of position of an axillary flower with respect to subtending bract
and the axis to which the bract pertains. The flower is placed
between the two. The portion of the flower which faces the
subtending bract is the anterior, likewise called inferior or lower.
The opposite portion which faces the axis of
inflorescence is the posterior, or superior, or
upper. The right and left sides are lateral.
( Fig< 804 ' 306 *) These relations do not
\ ? Appear in a solitary flower terminating a
/ simple stem ; but when such an axis produces
axillaiy branches with a terminal flower, the
relation of this flower to the preceding axis
and its leaf is manifest, just as in indetermi-
nate inflorescence.
291. Median and Transverse. The position of parts which lie
in antero-posterior line, or between bract and axis, is median.
Thus, in Fig. 304 and 305, the parts are all in the median plane :
in Fig. 306, the bractlets, b', b', are lateral or collateral, or (being
in the opposite plane) transverse.
292. Position of Bractlets. The rule has already been laid
down (285) that the first leaf of an alternate-leaved secondary
axis is in Monoctyledons usually median and posterior, that is,
farthest away from the subtending leaf (as in Fig. 304, 305) ;
in Dicotyledons, lateral or transverse. When these secondary
axes are one-flowered peduncles or pedicels, the leaf or leaves
(if an} T ) the}' bear are bractlets* Commonly there is only this
1 Not (with propriety, although the terms have been so used) exterior or
outer for the anterior, and interior or inner for the posterior position. These
terms should be reserved for the relative position on the axis of successive
circles or parts of circles, spirals, &c. Covering or overlapping parts are
exterior or outer in respect to those overlapped.
2 Latin Bracteohe : not that they are small bracts, but bracts of an ulti-
mate axis. In axillary inflorescence, the distinction between bractlet and
bract is obvious: in case of a solitary terminal flower, there is no ground of
difference : in terminal or cyrnose inflorescence, the difference is arbitrary ;
but we may restrict the term bractlet to the last bract or pair.
German botanists mostly distinguish between bracts, as a leaf subtending
a flower or cluster, and bractlets, by terming the former a Deckblatt, and the
FIG. 306. Diagram (cross section) of papilionaceous flower and its relation to axis
(a), bract (b), and bractlets (/, V).
THE CYMOSE TYPE. 161
posterior one to a simple axis in Monocot3 r ledons, and two
transverse ones in Dicot3 r ledons, i. r. one to the right and the
other to the left of the subtending bract, Fig. 306, If b'. When
the latter form a pair, they are perhaps always truly transverse ;
when alternate, the}' stand more or less on the opposite sides and
transverse. When more than one in Monocotyledons, they may
become either median or transverse, or even intermediate. The
relation of bractlets or bract, that is, of the last leaves of inflo-
rescence, to the first of the blossom, might be considered either
under Phyllotaxy or under Floral Symmetry. In general, it
may be noted that successive members stand over the widest
intervals ; l in other words, that the first leaf of the flower is as
far away as may be from the highest bractlet. For instance,
when there is a single and posterior bractlet, as is common in
Monocotyledons, the first leaf of the flower is anterior, the
next two right and left at 120. When there is a single and
lateral bractlet and five leaves in the first circle of the flower
(which occurs only in Dicotyledons), the first leaf of this circle
is either exactly on the opposite side from the bract, or at a
divergence of two fifths, the latter falling into the continuous
spiral. When with a pair of bractlets, right and left, the first
flower-leaf is at J divergence from one (the uppermost) of them
when the circle is of three, or at when of five members, or near
it ; but with many exceptions. 2
A tabular view of the kinds of inflorescence and their termi-
nology, serving as a key, may aid the student. 8
latter, being the leaves which the new axis first bears, Vorblatter, which is
also the name they apply to primordial leaves in germination.
1 In accordance with Hofmeister's law ; but (as Eichler remarks) not to
be explained on his mechanical principle of production in this place because
of the greater room : for the position of the first member of an axillary
flower is mostly the same as regards the subtending bract when the bractlets
are wanting.
2 When bractlets are wanting, the leaves of the first floral circle if two
are right and left ; if three, two lateral-posterior and one anterior ; when
five, the odd one commonly in the median line, either anterior or posterior.
3 INFLORESCENCE is either PURE, all of one type, or MIXED, of the two types
combined. The Types are :
I. Main axis not arrested and terminated by a flower. Indeterminate, Indefi-
nite, Acropetal or Ascendinr/, Centripetal, or BOTRYOSE.
II. Main and lateral axes arrested and terminated by a flower. Determinate,
Definite, Descending, Centrifugal, or CYMOSE.
.162 ANTHOTAXY, OK 1XFLOKESCENCE.
I. BOTRYOSE TYPE.
1. Simple, with lateral axes unbranched and terminated by a single flower, and
Flowers on pedicels,
Of somewhat equal length on a comparatively elongated axis RACEME.
The lower ones longer than the upper, and main axis short,
Of nearly equal length on an undeveloped main axis,
Flowers sessile on a very short main axis, . . . .
Flowers sessile on a comparatively elongated main axis, .
CORYMB.
UMBEL.
HEAD.
SPIKE.
A fleshy spike or head is a SPADIX.
A scaly-bracted spike is an AMENT or
CATKIN.
2. Compound, with lateral axes branched once or more, bearing clusters instead of
single flowers.
Irregularly racemosely or corymbosely compound, . . PANICLE.
Homogeneously and regularly compound, as
Racemes in a raceme, COMPOUND RACEME.
Corymbs corymbose, COMPOUND CORYMB.
Umbels in an umbel, COMPOUND UMBEL.
Spikes spicate, COMPOUND SPIKE.
Homogeneously compound, the secondary ramification
unlike the primary, as Heads racemose, Umbels spiked,
Spikes panicled, &c.
II. CYMOSE TYPE.
1. Simple, with terminal axis of each generation one-flowered.
Monopodial, the axis of each generation evidently re-
solved into branches, TRUE CYME.
These more than two, Pleiochnsium or MULTIPAKOUS CYME.
These only two, . . . Dichasium, Dich otomous or BIPAROTS CYME.
Sympodial, the apparently simple axis continued by
a succession of new axes standing end to end,
Monachasium, False Raceme or Spike, Botryose or UNIPAROUS CYME.
Flowers one-ranked on one side of rhachis, HKUCOIIJ UNIPAROUS CYME.
Flowers two-ranked on one side of rhachis, SCORPIOID UNIPAROUS CYME.
2. Compound, with terminal axes for one or more earlier
generations bearing a cyme instead of -a single
flower, Various sorts of COMPOUND CYME.
III. MIXED INFLORESCENCE.
1. Anomalous simple, with unbranched one-flowered lateral
axes (287), such as ... PARTLY KEVERSED SPIKES OR RACEMES.
2. Compound, of various combinations, of which there are
names for the subjoined :
Primary- inflorescence botryose, with axis elongating;
secondary cymose, THYRSUS.
Pair of such opposite cymes seemingly confluent round
the main axis, VEKTICILLASTER.
Panicle with some of the ramifications cymose, . . . MIXED PANICLE.
THE FLOWER. 163
CHAPTER VI.
THE FLO WEE.
SECTION I. ITS NATURE, PARTS, AND METAMORPHY.
293. FLOWER-BUDS are homologous with (morphologically an-
swering to) leaf-buds, and they occupy the same positions. (266.)
A FLOWER is a simple axis or a terminal portion of one, in a
phomogamous plant, with its leaves developed in special forms,
and subservient to sexual reproduction instead of vegetation.
294. In passing from vegetation to reproduction, it is not
always easy to determine exactly where the flower begins. The
same axis which bears a flower or floral organs at summit bears
vegetative leaves or foliage below. Or when it does not, as
when an axillary flower-stalk or pedicel is bractless, the change
to actual organs of reproduction is seldom abrupt. Usually
there are floral envelopes, within and under the protection of
which in the bud the essential organs of the flower are formed.
Some or all of these protecting parts, in very many flowers, are
either obvious leaves or sufficiently foliaceous to suggest their
leafy nature ; and even when the texture is delicate, and other
colors take the place of the sober green of vegetation, they are
still popularly said to be the leaves of the blossom. These pro-
tecting and often showy parts, though not themselves directly
subservient to reproduction, have always been accounted as
parts of the flower. 1 Between the lowest or outermost of these
and the bractlets and bracts there are various and sometimes
complete gradations. The axis itself occasionally undergoes
changes in such a way as to render the determination of the
actual beginning of the flower somewhat arbitrary. Moreover,
the flower itself is extremely various in different plants, in some
consisting of a great number of pieces, in others of few or only
one ; in some the constituent pieces are separate, in others
combined. The flower is best understood, therefore, by taking
some particular specimen or class of flowers as a representative
1 Indeed, the colored leaves, or envelopes in whatever form, essentially
were the flower in most of the ante-Linnaean definitions (that of Ludwig
excepted), as they are still mainly so in popular apprehension.
1G4
THE FLOWER.
or pattern, and especially some one which is both complete and
morphologically simple.
295. Such a flower consists of two kinds of organs, viz. the
Protecting Organs, leaves of the blossom, or floral envelopes,
which, when of two sets, are CALYX and COROLLA ; and the
Essential Reproductive Organs, which co-operate in the production
of seed, the STAMENS and PISTILS.
296. Floral Envelopes, Perianth, or Perigone, the floral leaves
or coverings. The former is a proper English designation of
these parts, taken collectively. But in descriptive botany,
where a single word is preferable, sometimes the name perianth
(Lat. pericmthium} , sometimes that of peri gone (or perigonium) ,
is used. PerianthiumJ- a Linnsean term, has been objected to,
because it etymologically denotes something around the flower ;
but it seems not inappropriate for the envelopes which surround
the essential part of the flower. Periyonium,
a later term, has the advantage of meaning
something around the reproductive organs,
which is precisely what it is. Neither name
is much used, except where the perianth or
perigone is simple or in one set (when it is
almost always calyx), or where it is of two
circles having the gen-
eral appearance of one
and needing descrip-
tive treatment as such,
as in the petaloideous
Monocotyledons. It is
also used where the
morphology is ambigu-
ous. Generally, the floral envelopes are treated distinctively as
calyx and corolla, one or the other of which (mostly the corolla)
may be wanting.
297. The Calyx is the outer set of floral envelopes. That is
its only definition. CommomV it is more herbaceous or foliaceous
than the corolla, and more persistent, yet sometimes, as in the
Poppy family, it is the more deciduous of the two. Not rarely it
1 Linnaeus (and about the same time Ludwig) used it in the sense of a
proper calyx, yet with some vagueness. Mirbel and Brown established it
in the sense of the collective floral covering. DeCandolle revived Ehrhart's
FIG. 307. The complete flower of a Crassula. 308. Diagram of its cross-section in
the bud, allowing the relative position of its parts The five pieces of the exterior
circle are sections of the sepals; the next, of the petals; the third, of the stamens
through their anthers; the innermost, of the live pistils.
FIG. 309 A sepal; 310, a petal; 311, a stamen; and 312, a pistil from the flower
represented in Fig. 307.
ITS NATURE AND PARTS. 165
is as highly colored. A name being wanted for the individual
leaves which make up the calyx, analogous to that for corolla-
leaves, DeCanclolle adopted Necker's coinage of the word sepal.
Calyx-leaves are SEPALS.
298. The Corolla is the inner set of floral envelopes, usually (but
hot always) of delicate texture and other than green color, form-
ing therefore the most showy part of the blossom. Its several
leaves are the PETALS. 1
299. The floral envelopes are for the protection of the organs
within, in the bud or sometimes afterward. Also, some of them,
by their bright colors, their fragrance, and their saccharine or
other secretions, serve for allurement of insects to the blossom, to
mutual advantage. (504.) This furnishes a reason for neutral
flowers, those devoid of essential organs, which sometimes
occur along with less conspicuous perfect ones. "The leaves of
the flower " are therefore indirectly subservient to reproduction.
300. The essential organs, being commonly plural
in number, sometimes need a collective name. Where-
fore, the aggregate stamens of a flower have been
called the ANDRCECIUM ; the pistils, the GYN cal section, and Fig. 315, by Linnaeus
named Germeri), the hollow portion
B at the base which contains the
OVULES, or bodies destinetl to be-
come seeds ; the STYLE (6), or colum-
nar prolongation of the apex of the
ovary ; and the STIGMA (c) , a portion
of the surface of the style denuded of epidermis,
sometimes a mere point or a small knob at the
apex of the style, but often forming a single or
double line running down a part of its inner face,
and assuming a great diversity of appearance in
different plants. The ovary and the stigma are the
essential parts. The style (as also the filament of a
stamen) may be altogether wanting.
apices. Ifc came in time to be used as now for the whole organ ; but Lud-
wig (Inst. Reg. Veg.), in 1742, apparently first so defined it, and introduced
the term Anther for the Apex of Ray, or Theca of Grew.
1 Following Linnaeus, this term is here freely used in the plural, and for
each actual separate member of the gynoecium, each organ which has an
ovary, stigma, and commonly a style. Tournefort, who appears to have
introduced the word, employed it in the sense of gynoecium. Many authors
define it thus, and then practically eliminate from botany this, one of the
oldest of its terms, and one by no means superfluous. The typical pistillum
of Tournefort is that of the Crown Imperial (Inst. i. 69, & tab. 1) and the
name is from the likeness to a pestle in a mortar. As it soon became im-
possible to apply the same name to the pistil of a Fritillaria or of a Plum,
the cluster of such organs in Caltha, and the capitate cluster and receptacle
of such organs in a Ranunculus or Anemone, Linnaeus, and Ludwig before
him, took the idea of Tournefort's name, and used it accordingly.
"Pistillum est pars interior et media floris, quae ex ovario et stylo com-
ponitur. . . . Ovarium est pars pistilli inferior, quae futuri fructus delinca-
tionem sistit. . . . Stylus est pars pistilli ex ovario centro producta. . . .
Summitas styli vel ejus partiurn Stic/ma dicitur." Ludwig, Inst. Reg. Veg.
41-43, 1742. Without mentioning the plural, the pistil is thus defined in a
way which necessitates its use. Linnaeus (in Phil. Bot.) first defines Stamen
and Pistillum in the singular number, enumerating the three parts of the
latter, and afterwards (p. 57) declares that "Pistilla differunt quoad
FIG. 314. Vertical section of a pistil, showing the interior of its ovary, a, to one
sMe of which are attached numerous ovules, d: above is the style, b, tipped by the
stigma, c.
FIG. 315. A pistil of Crassula, like that of Fig. 312, but more magnified, and cut
across through the ovary, to show its cell, and the ovules it contains; also pulled open
below at the suture. At the summit of the style is seen a somewhat papillose portion,
destitute of epidermis, extending a little way down the inner face : this is the stigma.
ITS NATURE AND PARTS.
167
303. The Torus, or Receptacle of the flower, also named
THALAMUS, 1 is the axis which bears all the other parts, that
upon which they are all \ interfere with floral symmetry, sometimes to
such degree that the true plan of the blossom
is to be ascertained only through extended
comparisons with the flowers of other plants
of the same order or tribe, or of related
orders. The symmetry of the blossom finds its explanation in
the laws which govern the arrangement of leaves on the axis ;
that is, in ph}ilotax3 T . The deviations from sj'mmet^ and from
t} T pical simplicit}' have to be explained, and in the first instance
1 For convenient reference and the avoidance of circumlocution, some
writers term the stamens which are before the petals ep//>eta/ows, those before
the sepals e/n'se/xi/ous ; but, as this prefix means upon, it is better to restrict
these terms to cases of adnation of stamens to these respective parts of the
perianth, and to distinguish as
^.n^'peto/oMS, those stamens which stand before petals, whether adnate or
free, and
.4??^'se/?af/oMS, those which stand before sepals. These terms we find have
already been employed in this way by Dr. A. Dickson (in Seemann, Jour.
Bot. iv. 275), with the addition of a third, viz.
Parapeta/ous, for stamens which stand at each side of a petal, yet not
necessarily before a sepal, as in many Rosaceae.
2 These particular types, with their modifications, are set forth in the
cfiarae/ers or distinguishing marks of the orders, tribes, genera, &c. The
best generally available illustrations of ordinal types are in Le Maout and
Decaisne's Traite Gene'ral de Botanique, and in Hooker's English edition
and revision, entitled A General System of Botany, Descriptive and Analyti-
cal, London, 1873. The best morphological presentation is in Eichler's
Bliithendiagramme, &c. (Flower Diagrams, Constructed and Illustrated),
Leipzig, 1875.
FIG. 330. Opened flower of Trillium erectum. 331. Diagram of the same.
ITS GENERAL MODIFICATIONS. 179
to be classified. To have morphological value, such explanation
should be based upon just analogies in the foliage and other
organs of vegetation. Whatever is true of leaves and of the
vegetating axis as to position of parts, mode of origin and
growth, division, connection, and the like, may well be true of
homologous organs in the flower.
SECTION III. VARIOUS MODIFICATIONS OF THE FLOWER.
1. ENUMERATION OF THE KINDS.
326. In the morphological study of flowers, these modifica-
tions are viewed as deviations from type. Their interpretation
forms no small part of the botanist's work. They may be classed
under the following heads :
1. Union of members of the same circle: COALESCENCE.
2. Union of contiguous parts of different circles : ADNATION.
3. Inequalit}' in size, shape, or union of members of the same
circle : IRREGULARITY.
4. Non-appearance of some parts which are supposed in the
type : ABORTION or SUPPRESSION.
5. Non-alternation of the members of contiguous circles :
ANTEPOSITION or SUPERPOSITION.
6. Increased number of organs, either of whole circles or
parts of circles : AUGMENTATION or MULTIPLICATION.
7. Outgrowths, mostly from the anterior or sometimes pos-
terior face of organs : ENATION.
8. Unusual development of the torus or flower-axis.
9. To which may be appended morphological modifications,
some referable to these heads and some not so, which are in
special relation to the act of fertilization. These are specially
considered in Section IV.
327. These deviations from assumed pattern are seldom single ;
possibly all may coexist in the same blossom. Several of them
occur even in that one of the. orders, the Crassulacese, which
most obviously exhibits the normal type throughout.
328. Thus, Sedum (Fig. 321)), with two circles of stamens,
being taken as the true t}'pe (324), Crassula (Fig. 307) wants
the circle of stamens before the petals ; Tillaea (Fig. 326) is the
same, but with the members sj-mmetricall}- reduced from five to
three ; Rhodiola loses all the stamens by abortion in one half
the individuals and the pistils in the other, sterile rudiments
testifying to the abortion ; Triactina has lost two of its five
carpels, and the three remaining coalesce into one body up to the
180
THE FLOWER.
middle ; Penthorum (Fig. 335, 336) has its five carpels coales-
cent almost to the top, and usually loses its petals by abortion ;
in Grammanthes and Cotyledon (Fig. 332-334), the sepals are
coalescent into a cup and the petals into a deeper one, out of
which the stamens appear to arise, these being adnate to the
corolla. Symmetrical increase in the number of members of
each circle is no proper deviation from t}^, at least in this
family (in which flowers on the same plant sometimes vary from
5-merous to 4-merous and-6-merous) ; and in Sempervivum (to
which Houseleek belongs) these members are always more than
five and sometimes as many as twenty in each circle.
2. REGULAR UNION OF SIMILAR PARTS.
329. Coalescence, or the cohesion by the contiguous margins
of parts of the same circle or constituent set of organs, is so fre-
quent that few flowers are completely free from it. The last
preceding figures show it in the gynoecium and corolla. Fig.
471-476 further illustrate it in the corolla, and in various degrees
up to entire union ; and Fig. 483-488 illustrate it in the andrce-
cium. The technical terms which coalescence calls for, and
which are needful in botanical description, may be found under
the account of the particular organ, and in the Glossary. Such
growing together of contiguous members in the blossom is strict!} 7
paralleled by connate-perfoliate leaves of ordinaiy foliage (212,
Fig. 215), where it more commonly occurs in upper leaves, and
in bracts, which are still nearer the flower.
330. It should now be hardly necessary to explain that the
terms coalescence, cohesion, union, and the corresponding phrases
PIG, 332. Flower of Grammantbes. 333. Flower of a Cotyledon. 334. The corolla
laid open showing the two rows of stamens inserted on it. 335. The five pistils of
Penthorum, united. 336. A cross-section of the same.
UNION OF PARTS. 181
in the next paragraph, do not mean that the parts were once sepa-
rate and have since united. That is true only of certain cases. The
union is mostly congenital, equally so in the disks of foliage of
the Honeysuckle (Fig. 215) and in the corolla of a Convolvulus.
The lobes which answer to the tips of the constituent leaves of
the cup or tube are usually first to appear in the forming bud,
the undivided basal portion comes to view later. It might be
more correct to say that the several leaves concerned have not
isolated themselves as they grew. Accordingly, Dr. Masters
would substitute for coalescence and adnate the term mseparate.
But the common language of morphology needs no change, as it
consistently proceeds on the idea, and the prevalent fact, that
leaves are separate things, and that the tube, cup, or " inscpa-
rate " base of a calyx or corolla, consists of a certain number of
these. It is no contradiction to this view that they developed
3. UNION OF DISSIMILAR on SUCCESSIVE PARTS.
33 1 . Actuation is the most appropriate term to denote the organic
and congenital cohesion or consolidation of different circles, the
1 If it were seriously proposed to change the language of descriptive
botany in this regard, consistency would require its total reconstruction, with
the abolition of all such terms as deft, parted, &c. ; for the structures in
question are no more cleft than they are united. While these convenient
and long-familiar terms are continued in use (as they surely will be), although
quite contrary to literal fact, it cannot be amiss to continue those, such as
connate, adnate, coalescent, &c., which imply and suggest the fundamental fact
in the structure of phaenogahious and the higher cryptogamous plants, viz.
that leaves are normally unconnected organs.
Whether fusion or separation is the more complex condition, and therefore
indicative of higher rank, is a question of a different order. It is argued
that the fusion or lack of separation is an arrest of development, and there-
fore an indication of low rank or less perfection than the contrary. But a
phylogenetic view of the whole case may reverse this conclusion as respects
the blossom. The course of development from thallus and frond to distinct
foliage on an axis, from little to full differentiation, is clearly a rise in
rank, as also is the differentiation of foliage into ordinary leaves, petals,
stamens, and pistils. But there is as much differentiation in the flower
of a Convolvulus as of a Ranunculus, and more in that of a Salvia, a
Lobelia, and an Orchis. In all such flowers, the combination, the irregu-
larity, and the diversification in many cases of the members of the same circle,
all indicate complexity, greater specialization, and therefore higher rank.
The production of leaves distinct from the axis is one step in the ascending
scale : such specializations and combinations of these as occur in flowers are
higher steps ; and the most specialized, complex, and therefore highest in
rank are complete, corolliferous, irregular flowers, with a definite number of
members, and these combined in view of the adaptations by which the ends
of fertilization and fructification are best subserved.
182
THE FLOWER.
apparent growing of one part on or out of another, as of
the corolla out of the calyx, the stamens out of the corolla, or
all of them out of the pistil. This disguises the real origin
of the floral organs from the receptacle or axis, in successive
series, one within or above the other. Organs in this condition
are also and rightly said to be Connote (bora united) ; but, as
this terai is equally applicable to the coalescence of members of
the same circle, the word Adnate is preferable, as applying to
the present case only. Adnation is heterogeneous organic co-
hesion or adhesion : coa-
lescence is homogeneous
cohesion or union.
332. Adnation occurs in
very various degrees, and
affects either some or all the
organs of the flower. Its
consideration introduces into
terminology several peculiar
terms, which ma}' here be
defined in advance. Three
of them, introduced and
prominently emplo3'ed by
Jussieu, depend upon the
degree of adnation, or the
absence of it, viz. :
Bypogynous (literally be-
neath pistil) , applied to parts
which are inserted (i. e. are
borne) on the receptacle of
the flower, as in Fig. 336.
This is the absence of
adnation, or the condition
which corresponds with the
unmodified t}*pe.
Periyynons (around the
pistil) implies an adnation
which carries up the inser-
tion of parts (which always means apparent origin or place of
attachment) to some distance above or away from the recep-
FIG. 336*. Vertical section of a flower of the Common Flax, showing the normal or
hypogynous insertion of parts upon the torus or receptacle.
FIG. 337. Vertical section of a flower of the Cherry, to show the perigynous insertion,
or adnation to the calyx, of the petals and stamens.
FIG. 338. Similar section of the flower of the Purslane, showing an atluation of all
the parts with the lower half of the ovary.
UNION OF DISSIMILAR PARTS.
183
tacle, so commonly placing this insertion around instead of
beneath the pistil ; whence the name. The perigyny ma}' be,
as the figures show, merely
the adnation of petals and
stamens to calyx, the calyx
remaining hypogynons, as in
Fig. 337 ; or else the adna-
tion of the calyx, involving
the other organs, to the lower
part of the ovary, as in Fig.
338, or up to the summit of
the ovary, while the petals
and stamens are adnate still further to the calyx, as in Fig. 339.
The latter passes into what is called
Epigyuoiis (on the pistil), where the adnation is complete to
the very top of the ovary, and none beyond it, as in Fig. 340,
341. Yet here the parts so termed are not really on the ovary,
except where an epigynotis disk (394) actually surmounts it.
333. Adnation brings some other terms into use in botanical
descriptions, especially those of superior and inferior. In this
connection, these words (in Latin taking the form of snperus and
infertis) denote the position in respect to each other of ovary and
floral envelopes, not the morphological, but the apparent posi-
tion or place of origin. Thus, in Fig. 336 and in 337, the calyx
is inferior, or in other words the ovary superior. Here real and
apparent origin agree, this being the normal condition, which
is otherwise expressed b}' saying that the parts are/ree, i. e. free
from all adnation of one to the other. But, in Fig. 339-341, the
FIG. 339. Similar section of a flower of Hawthorn, showing complete adnation to
the summit of the ovary and of the other parts beyond.
FIG. 340. Vertical section of a Cranberry-flower, and 341, of flower of Aralia
mulieatilis, with so-called epigynous insertion of calyx, corolla, and stamens; the calyx
of the latter completely consolidated with the surface of the ovary, or its limb
obsolete.
184 THE FLOWER.
ovary is said to be inferior and the calyx superior, the catyx
and other parts, in consequence of the actuation of its lower part,
seeming to rise from the summit of the ovary.
334. Adnation of floral envelopes to pistil rarely extends
beyond the ovary ; yet, in species of Iris having a tube to
the perianth, this tube is common!}- adnate for most of its length
to the style. But when the calyx has its tube or portion with
united sepals prolonged, the petals and the stamens are usually
adnate more or less to it, i. e. are inserted on the calyx. And,
when the petals are united and prolonged into a tube, the sta-
mens, being within the corolla, are commonly adnate to or
inserted upon this.
335. No one doubts that the view is a true one which repre-
sents the perianth-tube as adnate to the style in Iris, petals and
stamens as adnate to calyx in the Cherry (Fig. 337), stamens
as adnate to base of corolla in Fig. 334, and a long way farther
in Phlox, &c. That the calyx is similarly adnate to the ovary
is nearly demonstrable in certain cases.
336. But, as the lower portion of a pear is undoubtedly recep-
tacle, or rather the enlarged extremuy of the flower-stalk, as in a
rose at least a portion of the hip is receptacle, as the tube of the
flower in a Cereus or other Cactacea has all the external char-
acters and development of a branch, so it is most probable that
in many cases the supposed calyx-tube adnate to an inferior
ovary is partly or wholly a hollowed receptacle (in the manner
of a Fig-fruit) ; that is, a cup-shaped or goblet-shaped develop-
ment of the base of the floral axis. This would bring the case
under 7. (326, 495.)
4. IRREGULARITY OF SIMILAR PARTS.
337. Irregularity, or inequality in form or in union of mem-
bers of a circle, is extremely common, either with or without
numerical symmetiy. One or two examples may suffice.
338. Irregular flowers with symmetrical perfection, except in
the gynoecium, are well seen in the Pea Family, to which belongs
the kind of corolla called Papilionaceous, from some imagined
resemblance to a butterfly. (Fig. 342-344.) This flower is
5-merous throughout, has the full complement of stamens (10, or
two sets) , but the gynoecium reduced to a single simple pistil.
The striking irregularity is in the corolla, the petals of which
bear distinguishing names : the posterior and larger one, exter-
nal in the bud, is the VEXILLUM or STANDARD (Fig. 344, a) ;
INEQUALITY OF SIMILAR PARTS.
185
the two lateral next and under the standard, AL.E or WINGS (b) ;
the two anterior, covered by the wings and partly cohering to
345
form a prow-shaped body (c) , the CARINA or KEEL. The catyx
is slightly irregular by unequal union, the two upper sepals
united higher than the
other three. The sta-
mens are much more
coalescent,butwith an
irregularity, nine com-
bined by the lower part
of their filaments, and
one (the posterior)
separate. (Fig. 345.)
339. The plan and
floral symmetry in the Locust-blossom and
its relatives are little obscured by the irregu-
larities and the coalescence, hardly more so
than in the plainer flower of its relative,
Baptisia (Fig. 347, 348), in which the petals are somewhat
alike, and the ten stamens are distinct or unconnected. Only
the calyx is more irregular, b} r the union of the two posterior
sepals almost to the tip. (Fig. 348.)
FIG. 312. Diagram of flower of tlie Locust, Robinia Pseudacacia : a. axis of inflores-
cence; 6. bract; first, circle of 5, calyx; five remaining pieces, corolla; next anthers,
10 in number: in the centre a single simple pistil. 343 Front view of Locust-flower
showing only the corolla. 341. This corolla displayed.
FIG. 345. And reed um of the Locust, nine stamens coalescent, one distinct. 346
Same of a Lupine, all ten filaments coalescent below into a closed tube.
FIG. 347. Calyx and corolla of Baptisia australis 348. Same with petals fallen,
showing ten distinct stamens and tip of the style.
186
THE FLOWER.
340. But in a Lupine-blossom, of equally near relationship,
a casual observer might fail to recognize the very same type,
although disguised only by cohesions. For while the two pos-
terior sepals are united to the tip on one side of the blossom, the
three others are similarly united into
one body on the anterior side, giving
the appearance of two sepals instead
of five : in the corolla, the two keel-
petals are more strictl}' united into a
slender scythe-shaped or sickle-shaped
body ; so that the petals might with
the unwary pass for four: in the
andrcecium, the coalescence includes all
ten stamens (Fig. 346), which is an
approach to regularity.
341. The 5-merous symmetry of the
Violet-blossom is complete until the
gyncecium is reached (but with only
one circle of stamens) ; the main irregu-
larity of the perianth is in the anterior
petal, with its nectariferious sac at base
(Fig. 349-351) ; the two stamens near-
est this send into the sac curious appendages, which the other
three do not possess ; the gynoecium is composed of three car-
pels coalescent into one compound ovary in a
manner hereafter explained. In Antirrhinum
and Linaria (Fig. 480, 481), there is a similar
/ \'l 6 ( N m f \ ^ rre ^ u ^ ar ^y accompanying coalescence of the
\ V I vv\l/ J'j) petals, the anterior one being extended at base
\V\Vvx Ml it | n O a nectariferous sac or hollow spur. 1 The
flower of a Lobelia (Fig. 488) has the same
numerical plan and sj'mmetiy as that of Viola
(except that the gyncecium is dimerous) ; but
thte members are adnate below and coalescent above, and the
corolla is irregular through unequal coalescence of the five petals,
and the absence of coalescence down one side.
1 PELORIA is a name given by Linnaeus to an occasional monostrosity of
these floAvers (imitated in sundry others), in which the base of every petal, or
answering part of the corolla, is prolonged downward into a sac or spur.
The sac is, morphologically considered, a departure from normal regularity :
in the monster, symmetrical regularity is restored by the development of four
more sacs.
FIG. 349. Flower of Viola sagittata. 350. Its sepals and petals displayed. 351.
Diagram of a Violet-blossom, from Eichler, with bract orsubteivling leaf (below), a pair
of bractlets (lateral), and axis to which the subtending leaf belongs (above or posterior).
DISAPPEARANCE OF PARTS.
187
5. DISAPPEARANCE OR OBLITERATION OF PARTS.
342. Abortion or Suppression are somewhat s} T nonymous terms
to denote the obliteration or rather non-appearance of organs
which belong to the plan of the blossom. Abortion is applied
particularly and more properly to partial obliteration, as where
a stamen is reduced to a naked filament, or to a mere rudiment
or vestige, answering to a stamen and occupying the place of
one, but incapable of performing its office ; suppression, to abso-
lute non-appearance. Such vestiges or abortive organs justify
the use of these terms, the more so as all gradations are some-
times met with between the perfect organ and the functionless
rudiment which occupies its place. Such obliterations, whether
partial or complete, may affect either a whole circle of organs
or merely some of its members. The former interferes with the
completeness of a flower, and may obscure the normal order of
its parts. The latter directly interferes with the symmetry of
the blossom, and is commonly associated with irregularity.
343. Of parts of a Circle. Among papilionaceous flowers
(338) , different species of Erythrina have all the petals but one
(the vexillum, Fig. 344, a) much reduced in
size, in some concealed in the calyx, and in
every way to be ranked as abortive organs.
In Amorpha, of the same family, these four
petals are gone, leaving no trace, reducing
the corolla to a single petal. (Fig. 352, 35c>.)
This one is evidently the vexillum, both by
position and shape ; and the 5-merous type,
also the particular type of the family, are still
discernible in the five notches of the calyx,
the ten stamens, &c. In a related genus,
Parryella, even this last petal is wanting, and
the andrcecium is straight, all irregularity thus
disappearing through suppression.
344. Delphinium or Larkspur and Aconite or Monkshood
furnish good examples of flowers in which irregularity is accom-
panied by more or less abortion. The calyx of the Larkspur
(Fig. 354-356) is irregular by reason of the dissimilarity of the
five sepals, one of which, the uppermost and largest, is pro-
longed posteriori}' into a long and hollow spur. Within these,
and alternate with them as far as they go, are the petals, only
FIG. 352. Stamens and pistil of Amorpha fruticosa. 353. An entire flower of the
ante.
188
THE FLOWER.
four in number, and these of two shapes, the two upper ones
having long spurs which are received into the spur of the upper
sepal; the two lateral ones having a small but broad blade
raised on a stalk-like claw ; and the place which the fifth and
lower petal should occup}' (marked in the ground-plan, Fig. 356,
FIG 3">4. Flower of a Larkspur. 355. The five sepals (outer circle) and the four
petals (inner circle) displayed. 356. Ground-plan of the calyx and corolla.
FIG. 357. Flower of an Aconite or Monkshood. 358. The five sepals and the two
small and curiously shaped petals displayed ; also the stamens and pistils in the centre.
359. Ground-plan of the calyx and corolla; the dotted lines, as in Fig. 356, representing
the suppressed parts.
DISAPPEARANCE OF PARTS. 189
b}- a short dotted line) is vacant, this petal being suppressed,
thereb}^ rendering the blossom un symmetrical. In Aconite
(Fig. 357-359), the plan of the blossom is the same,
but the uppermost and largest of the five dissimilar
sepals forms a helmet-shaped or hood-like body ;
three of the petals are wanting altogether (their
places are shown by the dotted lines in the ground-
plan, Fig. 359) ; and the two upper ones, which ex-
tend under the hood, are so reduced in size and so
anomalous in shape that the}^ would not be recog-
nized as petals. One of these, enlarged, is exhibited
in Fig. 360. Petals and other parts of this and of va-
rious extraordinaiy forms were termed b}' Linnaeus
NECTARIES, a somewhat misleading name, as they
are no more devoted to the secretion of nectar than
ordinary petals or other parts are. In these flowers,
moreover, the stamens are much increased in number.
345. Analogous abortion of some of the stamens, along with
a particular iiregalaritj' of the perianth, especially of the corolla,
characterizes a series of natural orders with coalescent petals. 1
These flowers are all on the 5-merous plan (except that the
gynoecium is 2-merous), but with corolla, and not rarely the
calyx, irregular through unequal union in what is called the bila-
biate or two-lipped manner. The greater union is always median,
or anterior and posterior, and two of the coalescent members form
one lip, three the other. The two posterior petals form the
upper lip, the anterior and two lateral form the lower lip of the
corolla ; in the calyx, when that is bilabiate, this is of course
reversed. In some, as in Sage and Snapdragon, the bilabiation
of the corolla is striking (Fig. 479-481), and readily comparable
to the two jaws of an animal ; in others, the parts are almost regu-
lar. The suppression referred to is, in most of these cases, that
of the posterior of the five stamens, as in Fig. B61, where it is
complete. In Pentstemon (Fig. 362), a sterile filament regu-
larly occupies the place of the missing stamen. The position
sufficiently indicates its nature. This is also revealed by the
rare occurrence of an imperfect or of a perfect anther on this
1 These natural orders in which this occurs, or tends to occur, are the
Serophulariacea3 (Snapdragon, Pentstemon, Mimulus, c.), Orobanchaceie
(Beech-drops), Lentibulacese (Bladderwort), Gesneraceae (Gloxinia), Big-
noniaceae (Trumpet Creeper, Catalpa), Pedaliacese (Martynia), Acanthacese,
Labiatae (Salvia, Stachys), c.
FIG 360 A petal (nectary) of an Aconite, much enlarged.
190
THE FLOWER.
filament, a monstrosity, indeed, but the monstrosity is here
a return to normal S3'mmetry. The two stamens nearest the
suppressed or abortive one generally
share in the tendency to abortion, as
is shown by their lesser length or
smaller anthers : in the flower of
Catalpa, these two also are either im-
perfect or reduced to mere vestiges
(as in Fig. 363) : in very many other
plants of these families, even these
vestiges are not seen, and so the five
stamens are by abortion or complete
suppression reduced to two.
346. Suppression in the gynoecium
to a number less than the numerical
plan of the flower (as shown in the
perianth) is of more common occur-
rence than the typical number, and
the reduction is comparatively con-
stant throughout the genus or order.
A papilionaceous or other leguminous
flower with more than one or with all
five pistils is exceedingly rare, and
except in one pentacarpellary genus
is a monstrosity. Suppression of
the interior is more common than
of exterior organs. Want of room
in the bud may partially explain
this.
347. Suppression of whole Circles. Such suppression or rather
non-production in the actual blossom of whole series of organs
which belong to the type, and indeed are sometimes present in
that blossom's nearest relatives, is very common. It gives
occasion to several descriptive terms, which may be here defined
together. First, and in general, flowers are
Incomplete, in w r hich an^- one or more of the four kinds of organs
is wanting, whatever these may be ;
Apetalous, when the corolla or inner perianth is wanting ;
Monochlamydeous, where the perianth is simple instead of
FIG. 361. Corolla of Gerardia purpurea laid open, with the four stamens, the place
which the fifth should occupy indicated by a cross.
FIG. 3H2. Corolla of Penlstemon jrrandiHorus laid open, with its four stamens, and
a sterile filament in the place of the fifth stamen.
FIG. 3R3. Corolla of Catalpa laid oj>en, with two perfect stamens and the vestiges
of three abortive ones.
DISAPPEARANCE OF PARTS. 191
double, in which case the wanting set is generally (but not quite
always) the inner, or the corolla ;
Dichlamydeous, when both circles of the perianth (calyx and
corolla) are present ;
Achlamydeous, when both are wanting, as in Fig. 365. (These
three terms are seldom employed.)
Unisexual (also Diclinous or Separated) , when the suppression
is either of the stamens or the pistils. In contradistinction, a
flower which possesses both is Bisexual or Hermaphrodite.
Staminate, or Male, when the stamens are present and the
pistils absent ;
Pistillate, or Female, when the pistils are present and the
stamens absent ;
Moncecious (of one household), when stamens and pistils oc-
cupy different flowers on the same plant ;
Dicecious (of two households), when they occupy different
flowers on different plants ;
Polygamous, when the same species bears both unisexual and
bisexual or hermaphrodite flowers. This may occur in various
ways, from the greater or less abortion of either sex, either on
the same or on separate individual plants ; as Monceciously or
DioBciously Polygamous, according to the tendency to become either
monoecious or dio3cious. Recently Darwin has well distinguished
the case of
Gyno-dicecious, where the flowers on separate individuals are
some hermaphrodite and some female, but none male only ; and
Andro-dio2cious, of hermaphrodite flowers and male, but no
separate female. The latter is a less common case.
Neutral, as applied to a flower, denotes that both stamens and
pistils are wanting, a case neither rare nor inexplicable on
grounds of utility. (356, 504.)
Sterile and Fertile are more loosely used terms. A sterile
flower may mean one which fails to produce seed, as a sterile
stamen denotes one which produces no good pollen, and a
sterile pistil one which is incapable of seeding. But commonly a
sterile flower denotes a staminate one ; a fertile flower, one which
is pistillate, if not also hermaphrodite.
348. Suppressed Perianth. Almost universally, when the peri-
anth is reduced to a single circle, it is the inner, or corolla, which
is not produced. Or, rather, when there is only one circle or sort
of perianth-leaves, it is called calyx, whatever be the appearance,
texture, or color, unless it can somehow be shown that an outer
circle is suppressed. For since the calyx is frequently delicate
and petal-like (in botanical language, petaloid or colored, as in
192
THE FLOWER.
Clematis and Anemone, Fig. 364) , and the corolla is sometimes
greenish or leaf-like, the only real difference between the two is
that the calyx represents the outer
and the corolla the inner series.
Even this distinction becomes arbi-
trary when the perianth consists of
three or four circles, or of a less
definite number of spirally arranged
members.
349. Yet the only perianth obvi-
ously present may be corolla, as when
the calyx has its tube wholly adnate
to the ovary and its border or lobes obsolete or wanting. 1 Aralia
nudicaulis (Fig. 341) is an instance, likewise many Umbelliferge,
some species of Fedia or Valerianella, the fertile flowers of Nyssa,
and those Compositae which have no pappus. For PAPPUS,
the name originally given to thistle-down and the like, answers
to the border or lobes of a calyx attenuated and depauperated
down to mere fibres, bristles, or hairs. The name is ex-
tended to other and less obliterated forms. (644, Fig. 631-633.)
When the obliteration is complete, as in Mayweed (Fig. 630) ,
in some species of Coreopsis, &c., the corolla seems to be simply
continuous with the apex of the ovary. A comparison with
related forms reveals the real state of things. 2
350. So also in Hippuris, in which (along
with extreme numerical reduction of the other
floral circles) the calyx as well as corolla seems
to be wanting ; but the insertion of the stamen
on the ovary (epigynous) suggests an adnate
calyx, and near inspection detects its border.
351. Both calyx and corolla are really want-
ing in the otherwise complete and perfect (symmetrical and
1 In the flowers of the two common species of Prickly Ash (Zanthoxy-
lum) of the Atlantic United States, one has a double, the other a single
perianth (as shown in Gray, Gen. Illustr. ii. 148, t. 156) : the position of the
stamens gives a presumption that the missing circle of the latter is the calyx ;
yet it may be otherwise explained. In Santalaceaa there are some grounds
for suspecting that the simple perianth, although opposite the stamens, is
corolla ; and the foliaceous sepal-lobes of the female flowers of Buckleya
would confirm this, if these are true sepals rather than adnate bracts.
2 In the pappus of Compositae, every gradation is seen between undoubted
calyx, recognizable as such by structure as well as position, and diaphanous
scales, bristles, and mere hairs, wholly " trie-homes " as to structure, although
in the place of "phyllomes " and representing them.
FIG. 3f>4. Flower of Anemone Pennsylvania; apetalous. the calyx petalotl.
FIG. 365. Achlamydeous flower of Lizard's-tail (Saururus ceriums), magnified.
DISAPPEARANCE OF PARTS.
193
trimerous) flowers of Saurums, Fig. 365. But achlamydeous
blossoms are usually still further reduced to a single sex.
352. Suppression of one circle of stamens is of very common
occurrence. It is seen in different species of Flax ; which have
mostlv 5-merous perfectly symmetrical and complete flowers with
one set of stamens abortive. In some species (as in Fig. 367),
vestiges of the missing circle of stamens are conspicuous in
the form of abortive filaments, interposed between the perfect
stamens ; in others, these rudiments are inconspicuous or even
altogether wanting.
353. Suppressed Androecium or Gynoecium. This occurs with-
out or along with suppression in the perianth. In cases of the
former, vestiges of m 3n
the aborted organs
often remain to sig-
nify the exact nature
of the loss. Sepa-
ration of the sexes
(monoecious, diceci-
OMS, &c.) is the re-
sult of such suppres-
sion. In Menisper-
mum (Fig. 368, 369),
this is accompanied
by an actual doub-
ling of both calyx
and corolla. . The
dio3cious flowers of
Smilax are similarly
complete, except by
the abortion of one sex, but the calyx and corolla are single.
FIG. 366. Flower of a Lin um or Flax. 367. Androecium and gynoecium ; the former
of 5 perfect stamens, alternating with 5 rudiments of a second set.
FIG. 368, 369. Dioecious flowers of Moonseed, Menispermum Canadense: 368, Stami-
nate or male blossom ; 369, Pistillate or female, but with six abortive stamens, before
as many petals.
FIG. 370. A catkin of staminnte flowers of a Willow, Salix alba. 371. A single
staminate flower detached and enlarged (the bract turned from the eye). 372. A pistil-
late catkin of the same species. 373. A detached pistillate flower, magnified.
104
THE FLOWER.
354. Combined with suppression of Perianth. This, which is
found inmost amentaceous or catkin-bearing trees, in some with
partial suppression of perianth, is well illustrated
in Willows, the flowers of which are all uchlamydeous
and dioecious. (347.) The little scale (gland or nec-
tary) at the inside of each blossom might be sup-
posed to represent a perianth, reduced to a single
piece ; but an extended comparison of forms refers
it rather to the receptacle. Willow-blossoms (Fig.
370-372) are crowded in catkins, each one in the axil of a bract :
the staminate flowers consist of a few stamens merely, in this
species of only two, and the pistillate of a pistil merely. In
Salix purpurea, the male flower seems to be a single stamen
(Fig. 374) ; but it consists of two stamens, united into one
body. Here extreme
suppression is ac-
companied with co-
alescence of the
existing members.
355. Still more
simplified flowers,
but more difficult
to comprehend, are
those of Euphorbia,
or Spurge. These
are in fact monoeci-
ous ; and the female
flower is a pistil, the
male is a stamen.
The pistillate flower
(of three carpels,
their ovaries united
into one three-lobed
compound ovary)
surmounts a slender peduncle which terminates each branch of the
flowering plant. (Fig. 375.) From around the base of this pe-
duncle rise other smaller and shorter peduncles, each from the
axil of a slender bract, and surmounted by a single stamen,
which represents a male flower. (Fig. 376, 377.) This umbel-like
FIG. 374. A separate staminate flower of Salix purpnrea, with the stamens coa-
lescent (monadelphous and svngenesious). so as to appear like a. single one.
FIG. 375. Flowering branch of Euphorbia corollata. 37C. Calyx-like involucre
diviilecl lengthwise, showingthe staminate flowers around a pistillate flower (a). 377. A
more magnified staminate flower detached with its bract, a; its peduncle or pedicel b,
surmounted by the solitary stamen, c. 378. Pistil in fruit, cut across, showing the
three one-seeded carpels of which it is composed.
SUPERPOSITION OF SUCCESSIVE PARTS. 195
flower-cluster is surrounded and at first enclosed by an involucre
in the form of a cup, which imitates a calyx ; and the lobes of this
cup (the free tips of the calyx-leaves) in the present species are
bright white, so that they exactly imitate petals. Here, then, is
a whole cluster of extremely simplified flowers, taking on the
guise of and practically behaving like a single flower, the invo-
lucre serving as calyx and corolla ; the one-stamened male flowers
collectively imitating the androecium of a polyandrous blossom,
and surrounding a female flower which might pass for the pistil of
it. A series of related forms, from various parts of the world,
gives proof that this interpretation is the true one.
356. Suppression of both Andrrecium and Gynoecium. This
occurs in what are termed Neutral Flowers (347), such as are
conspicuous at the margin of the cymes of H}~drangea (Fig. 293)
and of Viburnum lantanoides and Opulus, also at the margin of
the head of flowers of Sunflower, Coreopsis (Fig. 287, 288), and
the like. In these and most other instances, the perianth of
which only the flower consists is much larger and more showy
than in the accompanying perfect flowers : in fact, their whole
utility to the plant, so far as known, is in this conspicuousness.
No plant normally bears neutral flowers only ; but in cultivation
all sometimes become so by monstrosity, as in the form of Vibur-
num Opulus called Snowball or Guelder Rose, also in "full
double " roses, pinks, &c. Occasionally flowers become sterile
and neutral b.y mere depauperation and abortion of perianth as well
as of essential organs, as in certain Grasses ; but such are
mostly vestiges of flowers rather than neutral blossoms.
6. INTERRUPTION OF NORMAL ALTERNATION.
357. Anteposition or Superposition is the opposition of succes-
sive (or apparently successive) whorls which normally alternate.
This result is brought about in different wa}'s, some of which are
obvious, while of some the explanation is hypothetical.
358. In the first place, there are cases of seeming anteposi-
tion, which are explained away on inspection. In a tulip, lily,
and the like, there is a perianth of six leaves and a stamen be-
fore each. The simple explanation is that the flower is not
6-merous, but 3-merous : there is a calyx of three sepals, colored
and mostly shaped like the three petals, which alternate with
these and are clearly anterior in the bud ; next, three stamens
alternate with the petals or inner circle of the perianth ; then
the three stamens of the inner circle, alternating with the preced-
ing, necessarily are opposite the three petals, as the first three are
196 THE FLOWER.
opposite the sepals. These organs altogether are in four whorls
of three, not in two of six members ; and the pistil at the centre,
of three combined members, is the fifth and final whorl.
359. The Barberry family exhibits a similar seeming ante-
position, which is more striking on account of a multiplication
of the members of the perianth. The calyx is of six sepals in
two circles, the corolla of six petals in two circles, the stamens
equally six ; and so each petal has a stamen before and a sepal
behind it. But, when properly viewed as a trimerous flower with
double circles of sepals and petals as well as of stamens, all is
S3*mmetrical and normal. Menispermum in the related Moonseed
family is in the same case, but the flower is trimerous, as seen
in Fig. 369 : in the male blossom this is obscured in the andrre-
cium (Fig. 368) by a multiplication of the stamens. 1 The
same thing occurs in the perianth and bracts of certain Clusiaceae,
in which the members counted as in fours are superposed, and
in some of which the double dimerous arrangement with apparent
anteposition extends through the corolla ; while, in other closel}'
related flowers, the corolla changes to simply tetramcrous and to
alternation with the preceding four sepals. This passes, in the
same family and in the allied Ternstroemiacese, into
360. Superposition by Spirals, as where five petals are ante-
posed to five sepals, b} r an evident continuation of pentastichous
phyllotax}* ; and the stamen-clusters of Gordonia Lasiantlius
are probably in this wa} r brought before the petals. 2 The flower
of Camellia is continuously on the spiral plan up to the gynoe-
cium ; but upon one which, from the bracts onward, rises from tho
to the | and f order or higher, throwing the petals of the rosette
in a full-double flower into numerous more or less conspicuous
vertical ranks.
361. Anteposition in the Androecmm. It is in the androecium
that real anteposition is most common, and also most difficult to
account for upon airy one principle. Doubtless it comes to pass
in more than one way. This condition is chiefly noticed when
the stamens are definite in number, and mainly in isostemonous
and diplostemonous flowers. (324.)
362. With Isostemony. Vitis (Fig. 379-381), also Rhamnus
(Fig. 4 IT), 416), and the whole Grape and Buckthorn families of
1 In Columbine (Aquiiegia), multiplication of the stamens in successively
alternating 5-merous whorls similarly brings the androecium into ten ranks;
so, when these stamens in double flowers are transformed into hollow-spurred
petals, these are set one into another in ten vertical ranks.
- Gen. Illustr. ii. 1. 140. But the petals alternate with the sepals in the
ordinary manner of the flower, though their strong quincuncial imbrication
suggests the spiral arrangement.
SUPERPOSITION OF SUCCESSIVE PARTS. 197
which the}- are the types, afford familiar cases of a single circle
of stamens placed before the petals. In Vitis, there are green
nectariferous lobes or processes from s sso
the receptacle, alternate with and inside
the stamens : there is no good reason to
suppose that they answer to a second
row of stamens. All isostemonous Por-
tulacacese have the stamens before the
petals ; and, when the stamens are fewer
than the petals, those which exist occupy
this position. Among the orders with
gamopetalous corolla, such anteposition
is universal in 1 lumbaginacese, Primu-
laceae, the related Myrsinacese, and in
most Sapotacese, in the latter usually
with some complications.
363. The earliest and the most obvious explanation of the
anomaly is that of the suppression of an outer circle of stamens,
and to this view recent morphologists are returning. 1 Observa-
tion supplies no vestige of proof of it in Rhamnaceae and Vitaceae ;
but, in the group of related orders to which the Primulaceae be-
long, evidence is not wanting. For Samolus and Steironema
both exhibit a series of rudimentary organs exactly in the place
of the wanting circle of stamens, which ma}' well be sterile fila-
ments. In the allied order Sapotaceae, while Chrysophyllum
has in these respects just the structure of Primulacese, and
Sideroxylon that of Samolus, Isonandra Gutta (the Gutta-percha
plant) has a circle of well-formed stamens in place of the sterile
rudiments of the preceding ; that is, alternate with the petals,
1 Eichler, Bliithendiagramme, passim, and in preface to Part II. xviii.,
relating chiefly to obdiplostemon y. The principal opposing view is that of
St. Hilaire, Duchartre, c., maintaining that corolla and stamens here repre-
sent one circle of organs doubled by median chorisis ; upon which see note
under a following paragraph. According to that hypothesis, there is no
androecial circle in such blossoms, or only vestiges of one, but the petals have
supplied the deficiency by a supernumerary production of their own! The
more plausible hypothesis of Braun, that of a suppressed interior circle
of extra petals, would restore the alternation, and make the extant sta-
mens the fourth floral circle, as does the adopted explanation. Braun's
hypothesis, if it insists that an extra row of petals is wanting, supposes the
suppression of that which very rarely exists ; but, if of stamens, then the
supposed suppression is of that which is so generally present, or with indi-
cations of presence, as properly to be accounted a part of the floral type.
FIG. 379. Flower of the Grape Vine, casting its petals before expansion 380. The
same, without the petals: both show the glands of the disk distinctly, within the
stamens* 381. Diagram of the flower.
198
THE FLOWER.
completing the symmetry of the blossom and the normal alterna-
tion of its members. This explanation of the anteposition of a
single circle of stamens is the more readily received, because it
well accords with the idea here adopted, that the androecium
of a typical flower should consist of two circles of stamens.
(324. ) The only serious objections to this explanation rise out of
the difficulty of applying it to analogous anteposition when both
circles are present.
364. For Diplostemony , the condition of two circles of sta-
mens, each of the same number as the petals, is also itself very
commonly attended by anteposition. In normal or Direct
Diplostemony, that which answers to the floral type com-
pletely, the antisepalous stamens (324, note) are the outer
and the antipetalous the inner series, and the carpels when
isomerous alternate with the latter and oppose the sepals ; the
alternation of whorls is therefore complete, as in the diagram,
Fig. 382. Such stamens, however, may actually occupy a single
line or coalesce into a
tube, without derange-
ment of the type. But
it as commonly occurs
that the antipetalous sta-
mens are more or less
exterior in insertion, and
then the carpels, when
isomerous, are alternate
with the inner and anti-
sepalous stamens, and therefore opposite the petals, as in the
diagram, Fig. 383. This arrangement takes the name of
Obdiplostemony. In it the normal alternation of successive
whorls is interrupted, so as to produce anteposition,
365. With Obdiplostemony. This condition prevails, more or
less evidently, in Ericaceae, Geraniaceae, Zygophyllaceae, Rutaeeae,
Saxifragaceae, Crassulaceae, Onagraceae, &c. (but in some of these
with exceptions of direct diplostcmony) ; also, accompanied
by a peculiar multiplication of members (380), in Malvaceae,
Sterculiaceae, and Tiliaceae. The explanation is difficult. The
Irypotheses may be reduced to three, neither of which is quite
satisfactory. There is, first, the hj-pothesis of St. Hilaire, ap-
plied to this as to the preceding case (to Rhamnus, Vitis, &c.),
that these exterior antipetalous stamens belong to the corolline
whorl ; in other words, that the petal and the stamen before it
FIG. 382. Diagram of pattern flower with direct diplostemony 383. Diagram
of similar flower with Obdiplostemony. Both from Euhler's Blutheiuliagramiue.
SUPERPOSITION OF SUCCESSIVE PARTS. 199
(whether adnate to or free from it) answer to one leaf which
has developed into two organs by a deduplication (372) taking
place transversely. This makes the inner and
antisepalons stamens the third floral circle
or the only truly andrcecial one, and sym-
metrically alternate with the petals on the one
hand and the carpels on the other. The
second hj'pothesis conceives that there is a
whorl suppressed between these antipetalous
stamens and the corolla : this, ideally restored,
gives symmetric succession and alternation to all the succeeding
whorls. The five glands in a Geranium-flower, alternate with and
next succeeding the petals (Fig. 384), were plausibly supposed to
represent this missing whorl, which according to Braun should be
an inner corolla ; according to others rather a primary circle of
stamens. The third is the recent hypothesis of Celakowsk}-,
which Eichler adopts : this regards the antipetalous stamens as
really the inner or second circle, and conceives that in the course
of development it has become external by displacement. The
difficulties of this Irypothesis are, first to account for this dis-
placement, and then for the anteposition of the carpels to the
assumed inner stamens in the great majority of these cases. 1
1 In the first part of the Bliithendiagramme, Eichler inclined to the first
hypothesis, that of St. Hilaire (now very much abandoned on accoun^ of
the feeble evidence that there is any such thing as transverse or median
chorisis); in the second, he discards this in favor of Celakowsky's view
(published in Regensburg Flora, 1875). As to members which are morpho-
logically interior becoming exterior by outward displacement, Eichler cites
the staminodia or sterile stamen-clusters of Parnassia (Fig. 400, 401), and
the corresponding antipetalous stamens of Limnanthes, as clearly interior
in the early flower-bud, but exterior at a later period ; states that the vascu-
lar bundles which enter these stamens generally are either inner as respects
those of the episepalous stamens or in line with them ; that in some cases
(as in many Caryophyilaceae) the real insertion of the stamens is that of
direct diplostemony, while the upper part of their filaments and the anthers
are external to the episepalous series; that in most families with obdiplos-
temony examples of direct diplostemony occur, and still more cases with both
stamineal circles inserted in the same line ; and that, as a rule, the episep-
alous stamens are either later or not earlier formed th:in the epipetalous.
As to the position of the carpels before antipetalous stamens and petals,
Celakowsky suggests that this may result from the outward recession of
those stamens affording more room there, while in the normal case the
greater space is over the episepalous stamens. And, indeed, exceptions
to the prevalent position are not uncommon both in direct diplostemony
FIG. 384. Diagram (cross- sect ion) of the flower of Geranium macnlatum, exhibiting
the relative position of parts, and the symmetrical alternation of circles, i.e. sepals,
petals, greenish bodies called glands, antipetalous stamens, antisepalous stamens,
carpels.
200 THE FLOWER.
366. The case of stamens in a cluster before the petals is a
complication of either of the foregoing with a peculiar kind of
multiplication, termed deduplication or chorisis. (372.)
7. IXCREASED NUMBER OF PARTS.
367. Augmentation in the number of floral members is one
of the commonest modifications of the type. It occurs in two
ways : 1st, by an increased number of circles or turns of spirals
in the flower, which is Regular Multiplication ; 2d, by the pro-
duction of two or three or of many organs in the normal place
of one, Chorisis or Deduplication. The first does not alter the
normal symmetiy of the blossom, although it may render it dif-
ficult or impossible to trace or demonstrate it. The second
apparently disturbs, or at least disguises, floral symmetry.
Either may be definite, or of a constant and comparatively
small number ; or indefinite, when too numerous for ready
counting, or inconstant, as the higher numbers are apt to be.
368. Regular Multiplication, or Augmentation of floral circles
or spirals, may affect any or all the four organs, but most com-
monly the andro3cium. When the perianth is much increased
in the number of its members, the distinction between catyx and
corolla, or even between bracts and corolla, is apt to disappear,
as* in most Cactaceous flowers (Fig. 317), Nelumbium, Caly can-
thus, &c. In these and similar cases, the members of the perianth
are prone to take a spiral instead of cyclic arrangement ; and this
and in obdiplostemony. Along with the lack of clear analogy to support
St. Hilaire's hypothesis of transverse deduplication, the similar orientation
of the vascular bundles in the petal and the stamen before it must, as
Celakowsky insists, be good evidence that these represent independent
leaves, and not superposed portions of one.
The main objection to the second hypothesis (that of a suppressed
/circle outside of the antipetalous stamens) is that this missing circle,
whether of petals or stamens, is not actually met with in any nearly re-
lated forms (for in Monsonia the fifteen stamens are otherwise explained) ;
also that there are transitions, as above mentioned, between obdiplostemony
and direct diplostemony. To Braun's theory that the glands behind the
antisepalous stamens in true GeraniaeeaB answer to suppressed phylla,
Eichler objects that these are present behind all ten stamens in Oxalideas;
also that all are wanting when the office of nectar-secretion, which they sub-
serve, is undertaken by some other part of the floAver, as by the calyx-spur
in Pelargonium and Tropaeolum. The first objection is forcible : the second
mixes morphological considerations with functional, and is inconclusive.
Abortive organs, preserved for their utility as nectaries, might totally dis-
appear when rendered useless by a different provision for the same function.
INCREASED NUMBER OF PARTS. 201
is even more true of greatly multiplied stamens and pistils, as
in Magnolia and Liriodendron, most Anonaceae, Ranunculus,
Anemone, and the like. But in Aquilegia, where the number five
is fixed in the perianth, the cyclic arrangement with alternation
of whorls prevails throughout
369. The definite augmentation of calyx and corolla b} r the
production of one additional whorl of each, and the seeming
anteposition which comes of it when the androecium remains
simply diplostemonous (in the manner of the Berberidaeeae,
Menispermaceae, &c., 359) has already been explained.
370. Similar increase to two whorls affecting the corolla only
characterizes Anonaceae, Magnoliaceae, Papaveracese, and Fuma-
riaceae. In all but the last order, this is accompanied by indefi-
nitely multiplied stamens, and mostly by an increased number of
carpels. In Fumariaceae, which has dimerous flowers, there is a
diminution by the suppression in most cases of half the normal
andro2cium, and also an augmentation of the other half by
chorisis. (372.)
371. Parapetalous Multiplication. Under this head may be
described an anomalous arrangement of augmented stamens
which prevails in the order Rosaceae, but is not peculiar to it. 1
The simplest case, but a rare one, is seen in the 10-stamened vari-
ety of some Hawthorns, as occasionally in Crataegus coccinea and
Crus-galli. The ten are in one circle and in pairs, the pairs
alternate with the petals. Some would say the pairs are before
the petals ; but the space between two stamens before each petal
is mostly rather wider than in the pair taken the other way.
The next case in order, as in 15-stamened Hawthorns, and
constantly in Nuttallia, adds to the above a simple interior circle
of five stamens, one directly before the middle of each petal.
Next, as in most Pomeae and many Potentilleae, there are twenty
stamens, thus placed, but with an additional circle of five alter-
nating with the preceding one. Next there are 25 in three
circles, the second circle as well as the first having ten stamens ;
and finally there are from 30 to 50, all probably in circles of ten
each. There is little doubt that the circles develop in centri-
petal order ; the inner successively the later. 2
1 It was first clearly described by Dr. A. Dickson, in Trans. Bot. Soc.
Edinb. viii. 468, and Seemann's Jour. Bot. iv. 473 (1866). He introduced the
term, parapetalous, which is characteristic of it in its elementary form (254,
note) : it is particularly illustrated by Eiehler, in Bluthendiagramme, ii.
495-510. The former interprets it by chorisis, both median and collateral :
the latter presents the facts and possible views, but declines to adopt either
of them.
2 Accordingly, the whole is probably to be explained by some modifica-
202 THE FLOWER.
372. Chorisis or Deduplication. Both these terms, and the
ideas which they denote, originated with Dimal, but were first
expounded b} r Moquin-Tandon. 1 The first word is Greek for a
separating or separation. The second is a translation of Dunal's
French word dedoublement (literall}* undoubling) , the ambiguity
of which, and of the original presentation of the case, long
retarded the right apprehension of the subject. Direnrption has
been suggested (by St. Hilaire) as a proper term. The mean-
ing simply is, the division of that which is morphologically
one organ into two or more (a division which is of course
congenital) , so that two or more organs occupy the position of
one. As thus used, chorisis is restricted, or nearly so, to the
homologues of leaves in the flower, and mainly to stamens and
carpels ; the division or splitting up of a petal or a sepal, when it
occurs, being expressed in the phrases which are applied to leaves.
Yet a compound leaf, especially one of the palmate type, is
a good tj-pe of chorisis, the several blades of a compound leaf
answering to the single blade of a simple leaf. It has been ob-
jected against the terms chorisis and deduplication that they
assume the division of that which has never been united ; but
so equally does the established terminology of foliage. A di-
vided leaf has never been entire.
373. Chorisis is complete when the parts concerned are dis-
tinct or separate to the ver} r insertion, as in the stamen-clusters of
Hypericum. The foliar form of this would be represented by
tion of the augmentation of circles. Dickson's hypothesis, that the two,
three, or five stamens which are more or less in face of each petal are all
deduplications of that petal, would come to be noticed under the next head,
but it may be dismissed at once. Yet that the pairs in the outer circle
represent each an antisepalous stamen, divided by chorisis (sometimes
incompletely) and much separated, is not improbable. The other tenable
explanation (which may be harmonized with the last) is that the outer
circle of stamens here rightly consists of ten members, respectively alternat-
ing with the sepals and petals taken as a whole. This makes them para-
petalous, and at the same time brings them under Hofmeister's general law
that new organs originate over intervals of those preceding, in this case over
the ten perianth-intervals directly. It also accords with Hartog's elucidation
of the accessory parts in the flower of Sapotaceae (in Trimen's Jour. Bot. 1878).
The inner circles are there sometimes 5-merous after the primitive type,
sometimes 10-merous in regular alternation to the preceding circles.
1 Moquin-Tandon, Essai des Dedoublemens, &c., Montpellier, 1826; Con-
siderations sur les Irregularite's de la Corolle, &c., in Ann. Sci. Nat. xxvii.
287, 1832; Teratologie Ve'ge'tale, 337. Dunal, Essai sur les Vacciniees,
1819, cited by Moquin (some pages printed, but never published) ; Conside-
rations sur la Nature et les Rapports de quelques-uns des Organes de la
Fleur, 1829. The next botanist to develop it was St. Hilaire, Morphologic
Ve'ge'tale, 1841.
CHORISIS OR DEDUPLICATION.
203
such sessile palmately compound leaves as those of some species
of Aspalathus. It is incomplete when division does not extend
to the base; as in Fig. 387, 393. Compare, as a
proximate homologue of this, a petal of Mignonette,
Fig. 385. But proper chorisis requires that the
supernumerar} T organs should be developed like
unto the original organ which is thus multiplied, or
should complete their sj-mmttry, whatever it be.
374. St. Hilaire distinguished two kinds of deduplication ; viz.,
collateral when the members stand side b}' side, and parallel
when an organ becomes double or multiple antero-posteriorly.
The latter, sometimes called vertical, and sometimes transverse,
is better named median chorisis. The collateral is the origi-
nal and typical chorisis. Most botanists incline to restrict the
name to this, and to give some other explanation and name to
the median form of augmentation. But some cases, such as
those of Tilia and Sparmannia, are clearly of the same nature
as the collateral, and may be a disguised form of it ; there are
others which may be explained in accordance with it ; and there
are such transitions between some of these and coronal out-
growths that the term chorisis is most conveniently made to
comprise augmentation or doubling in either plane. Distinct
anteposition, however, may be explained in other wa}*s. (357.)
375. Typical or Collateral Chorisis, in which the members,
together answering to one leaf, normally stand side by side,
occurs in many families of plants, and
in a variet}' of forms. A few are here
presented.
376. Elodes Virginica (a common
marsh plant of the Hypericum family) ,
like most of its near relatives, has its
calyx and corolla on the plan of five,
its stamens and carpels on the plan of
three, as is shown in the diagram, Fig. 386. This makes a break
in the s^ymmetry between the corolla and the stamens ; but all
within is in regular alternation when the three stamens of each
cluster are counted as one as their union at base into a ph
four petals, of somewhat various overlapping in aestivation,
which essentially alternate with the two single stamens and the
two pairs ; lastly, four sepals, alternating with the four petals as
a whole, the anterior and posterior overlapping the lateral ones
in the bud: Now the median (i. e. the anterior and posterior)
pairs of stamens occasionally have their contiguous filaments
conjoined, as in Fig. 397. If this were at all constant, the
inference would undoubtedly be that the case is one of chorisis,
and that the flower as to its essential organs is dimerous. This
is apparently the best explanation to be given. It assumes that
the chorisis is normally complete in the androecium of Cruciferae,
instead of incomplete, as in Fumariaceae. 1 And this view is
confirmed by the fact that the median stamens are simple and
307
1 The hypothesis here adopted, as to the androeeium, is that of Steinheil
(1839), and of Eichler (in Flora, 1865, 1872, and Bliithend. ii. 200), replacing
that of Kunth, 1833, &c., employed in former editions. The rejected view
makes the flower 4-merous up to the pistil, and the stamens all of one circle,
alternating with the four petals, the median stamens (as in our view) doubled
by chorisis. Krause and Wretschko (cited as above by Eichler) would
have the floral circles 2-merous and 4-merous by turns ; the calyx of two
2-rnerous circles (which it plainly is); the corolla of one 4-merous circle
FIG. 394. A cruciferous flower. 395. Diagram of such a flower, with position of
axis marked above it. 396. Tetradynamous stamens and the pistil. 397. A common
monstrosity of the same, two of the four inner stamens combined into a common
2-antheriferous body.
CHORISIS OK DEDUPLICATION. 207
single in Senebiera and many species of Lepidium, in which the
lateral or short stamens are at the same time abortive.
379. It is quite possible that chorisis may be extended to the
corolla of the cruciferous flower, and reduce the whole to a
symmetrical 2-merous plan, and to congruity in the perianth
also with Fumariaceae. The only obstacle is in the petals form-
ing a whorl of four where all the rest is 2-merous, for the sepals
are manifestly two decussating pairs. Now the median petals
of Hypecoum are deeply 3-lobed. An abortion of their middle
lobe would leave them almost two-parted : a little more would
separate them ; then they would imitate the four cruciferous petals
as in the diagram, Fig. 395. Applying this view to Cruciferae,
the blossom in the two orders would accord in having a 2-merous
three- whorled perianth, the first and third whorls median ; l as
also in the dimerous androecium, the first whorl of which is
lateral. The difference is that in Fumariaceae the two members
of the first whorl of stamens augment by chorisis into three, and
the second is wanting, or is present only in Hypecoum ; while in
Cruciferse the first whorl is simple (of the two short stamens) , and
the second is doubled. In Fumariaceae only the first whorl of the
perianth counts as calyx, and
the corolla is of two whorls ;
in Cruciferae, the first and
second whorls are calyx, the
inner sepals answering to the
outer petals of Fumariaceae.
380. Chorisis along with
anteposition of stamens is well
seen in Tilia or Linden, at least in the American species. In
these the indefinitely numerous stamens are in five clusters, one
before each petal (Fig. 398, 399), and there is a petal-like body
alternating with the calyx-members as a whole ; the short stamens following
as a 2-merous circle ; then the long stamens as a 4-merous circle ; lastly the
2-merous gynoecium. G. Henslow (in Trans. Linn. Soc. ser. 2, i. 105) would
have the flower 4-merous by the suppression of the fifth members of a
5-merous type, and a further suppression of half of the remaining exterior
stamen-circle, &c. Finally, there is the much better-maintained view that
the cruciferous flower is 2-merous throughout, as explained in the following
paragraph, 379.
1 This view was taken by Steinheil, ; n Ann. Sci. Nat. ser. 2, 337 (1839),
and is essentially reproduced by a Russian botanist, Meschajeff, in Bull.
Soc. Imp. Nat. Mosc. 1872.
FIG. 398. Diagram of the flower of Tilia Americana, the common American Lin-
den or Basswood.
FIG. 399. A detached stamen-cluster with its petal-like scale.
208
THE FLOWER.
in each cluster with which the stamens cohere. The explanation
by chorisis is that each cluster, petal-like body included, is a
multiplication of one stamen. The diagram (Fig. 398) accu-
rately shows that most of the stamens originate from the outer
side of the base of the petal-like portion : this is most naturally
explained by median chorisis. The superposition of the clusters
to the petals will take the same explanation as that of Rhamnus,
Vitis, &c. (Fig. 363.) That the androecium is here composed
of the inner circle merei3 T is partly confirmed by the alternation
of the carpels with the clusters. According to Duchartre, 1 the
development of the androecium in a Mallow indicates a similar
structure ; for the whole united mass originates from five protu-
berances, one before each forming petal and connected with it,
this by collateral chorisis forming a cluster of stamens, and the
five clusters coalescing as they develop into a tube of filaments,
such as in Fig. 485. Now Hibiscus and its near relatives have
a naked tip to the stamen-tube, ending usually in five teeth ;
and Sidalcea, as is most strikingly shown in the California!!
S. diploscypha, has two series of stamens, the outer (answering
to those of Malva and its relatives) in five membranaceous pha-
langes, superposed to the petals ; the rather numerous inner
series, more or less in phalanges, surmounts an interior filament-
tube. Whence it is inferred
that these, and the five teeth
terminating the column in
Hibiscus, represent the in-
ner stamineal circle which is
wanting in Malva, as it is in
Tilia. 2
381. The case of Parnas-
sia would be explained as
analogous to that of Tilia,
but with the stamen-clusters before the petals wholly sterile,
and of fewer divisions, while an inner circle of five stamens
1 Comptes Rendus, 1844, & Ann. Sci. Nat. ser. 3, iv. 123. Duchartre and
others who draw freely upon median chorisis to explain anteposition, and
consider that congenital union proves it, take the phalanges in these cases,
like the single stamens in Vitis, to be an inner part of the petal itself. But
this view appears to have had its day.
2 Gray, Gen. Illustr. ii. 44, 57, 75-82. The position of the carpels before
the petals in Pavonia and Malvaviscus brings the former into symmetrical
alternation with such an inner stamen-circle; but it is not so in Hibiscus,
which has the carpels before the sepals.
FIG. 400.
FIG. 401.
A petal of Parnnssia Caroliniana. with a triple staminodium before it.
Diagram of the flower of Parnassia Caroliniana.
OUTGROWTHS. 209
alternate with the petals forms the effective anclroecium. For
the scale-like body before each petal, and even slightly adnate
to its base (in P. Caroliniana about 3-parted, as in Fig. 400, but
in P. palustris a thin scale, fringed with more numerous gland-
tipped filaments), is plainly outside the stamens in the full-grown
flower-bud. But Eichler and Drude have found
that it is inside in the early bud. 1 Wherefore, if
these stamen-like bodies really represent a circle
of the andrcecium, it must be the inner one ; and
that is the more probable view.
382. Multiplication by chorisis in the gynoecium
is not common ; but there are well marked in-
stances of it in all degrees. In Drosera, the
styles and stigmas are doubled (Fig. 402) ; in
Malvaceae, the same thing takes place in Pavonia
and its allies ; while in Malope and two other
genera of the same order the few normal carpels are multiplied,
evidently by chorisis, into an indefinite number of wholly distinct
ones.
8. OUTGROWTHS.
383. Proper chorisis is the congenital multiplication of one
organ into two or more of the same nature and office ; or at
least into two or more organs, even if dissimilar, as in the
American Lindens, in which one member of the cluster is a kind
of petal. Between this and the production by an organ of ap-
pendages, or outgrowths of little or no morphological signifi-
cation, there are many gradations ; as also between these and
mere cellular outgrowths from the surface, even down to
bristles and hairs. The latter, in all their variety and modifica-
tions, are properly outgrowths of the epidermis only, and there-
fore consist of extended cells, single or combined, unaccompanied
by vascular or woody tissue. To them has been given the
general name of Trichomes (Trichoma, pi. trichomata)', that is
structures of which hairs are the t}-pe. They may occur upon
the surface of any organ whatever. Their morphology is the
morphology of cells rather than of organs. They will therefore
be most conveniently illustrated under Vegetable Anatomy as
1 Eichler in Fl. Brasil., Sauvagesiacse, & Bliithend. ii. 424 ; Drude in Lin-
naea, xxxix. 239. Eichler refers to this as a confirmation of Celakowsky's
explanation of obdiplostemony by posterior displacement. (365.)
FIG. 402. Pistil of Drosera filiformis with tricarpellary ovary (transversely divided),
and six styles, i. e. three, and each two- parted.
210
THE FLOWER.
respects their structure, and in the Glossary as respects ter-
minology.
384. But into some bristles, such as those of Drosera. a sub-
jacent stratum of tissue enters, including one or more ducts or
even some woody tissue. Prickles are of this class ; and from
the most slender, which pass into bristles, there are all grada-
tions of stoutness and induration. Such outgrowths ma} 7 even be
formed in most regular order, as the prickles on the calyx-tube
of Agrimonia and scales on the acorn-cup of Oaks, and yet have
no morphological importance. On the other hand, true represen-
tatives of leaf or stem may, by abortion and depauperation, be
reduced to the structure as well as the appearance of trichomes.
Examples of this are familiar in the pappus (answering to limb
of the calyx) of many Compositae, and in the bristles which
answer to perianth in many C3 7 perace3e. The scarious stipules
of Paronychia and of Potamogeton, the ligule of Grasses, and
even the corolla in Plantago, are equally reduced to mere cellular
tissue. So that the structural difference between trichomes and
outgrowths 1 is not at all absolute, and the morphological distinc-
tion must rest upon other ground than anatomical structure.
385. Among the corolline outgrowths most akin to chorisis is
the Crown ( Corona) of Silene and allied Caryophyllacese, at the
junction of the claw
with the blade of
the petals (Fig. 403),
the analogy and
probable homology
of which to the ligule
of Grasses (Fig.
1 50) is evident ; also
the many-rayed fila-
mentous crown of
Passion-flowers (Fig. 404) , which consists of two or more series
of such outgrowths. In Sapindus and some other Sapindaceae,
these ligular outgrowths or internal appendages are more like
a doubling of the petal ; as also in Erythroxylum, where they
1 This is the best English name for the Emergenzen of the Germans, the
Epiblastema of Warming, &c. For the development and discussion of this
subject, see Warming, in Kjobenhavn Vidensk. Meddel. 1872, and a larger
treatise on Ramification in Phanerogams, Copenhagen, 1872. Also, Uhl-
worm in Bot. Zeit. 1873 ; Celakowsky in Flora, 1874 ; and Eichler's note on
Emergenzen in Bluthendiagramme, i. 48.
FIG. 403. Petal of Silene Pennsylvania, with its crovm.
FIG. 404. Flower of Passiflura cserulea, reduced in size.
403
FORMS OF THE RECEPTACLE.
211
are often more complicated in structure. They are alwa}"S on
the inner face, and are commonly two-lobed or parted.
386. Similar stamineal appendages are well known in Cuscuta
(Dodder), in Larrea (Fig. 405) and other Zj'gophyl-
Iacea3, and less conspicuously in Gaura.
387. To extend to them the name of LIGTJLE may
not be amiss, whether they are regarded as mere
outgrowths of floral leaves, without further morpho-
logical relations, or wiiether they be, at least some-
times, interpreted as the homologue of intrapetiolar
stipules, as their ordinarily two-cleft form, and their
coincidence in Erythroxj'lum with an intrapetiolar two-cleft stipule
suggest.
9. FORMS OF THE TORUS OR RECEPTACLE.
388. Torus is the more specific and proper name, RECEPTACLE
is the more usual. (303.) A normal receptacle of the flower
would be that of Fig. 316, the apex of the flower-stalk somewhat
enlarged, roundish or depressed, and with surface mainly cov-
ered by the insertion of the several organs ; the several inter-
nodes which it potentially contains being
undeveloped. As the members of the flower
multiply and occupy numerous ranks, the
receptacle enlarges or lengthens to give them
insertion or standing-room.
389. Of elongated forms of receptacle,
Magnolia and Liriodendron or Tulip-tree give
familiar instances. The lengthening in the
former is mainly for the support of both an-
drcecium and gynoacium ; in the latter, as in
Myostirus, mainly for the gynoecium only.
The fall of the matured carpels reveals it
as a very slender or bodkin-shaped pro-
longed axis. Of broadened forms, the Straw-
berry, even in blossom, affords a familiar
example. (Fig. 406.) In the same order,
Rubus odoratus shows a very broad and flat
receptacle : in roses, it is so deeply concave as to become the
reverse of the strawbeny (Fig. 407), being urn-shaped with a
narrow mouth, upon which the petals and stamens are borne,
FIG. 405. Stamen of Larrea Mexicana, with a conspicuous ligulate appendage at
the base within.
FIG. 406. Receptacle of a strawberry in longitudinal section. 407. Same of a rose,
in diagram.
212
THE FLOWER.
while the pistils line the walls of the cavity, the base or centre
of this cavity answering to the apex of the strawberry.
390. Sometimes internodes are lengthened between certain
members. In Schizandra, the receptacle, barely oblong in blos-
som, lengthens greatly in fruiting, so as to scatter the carpels
on a long filiform axis.
391. In many Gentians, in Stanleya and Warea among
Cruciferae, and in most species of Cleome, the internode of the
receptacle between stamens
and pistil is developed into
;r,\ \\\\WJJ \ \ \ $7 // a lon S stalk to the latter -
l( \\\n/// Gynandropsis (Fig. 409) is
like its near relative, Cleome,
except that this very long stalk
- * ias *ke l wer P art f tne
iw/l \XilA stamens adnate to it : the in-
ternode between the corolla
and calyx is broad and slightly
elevated (or in Cleome, &c.,
narrower and longer) ; and
409 so the several floral circles
are as it were spaced apart by this unusual development of
receptacular internodes. In Silene (Fig. 408) and many other
plants of the Pink family, an internode between the calyx and
corolla is prolonged into a stalk or Stipe. 1
1 STIPE is the general name of a stalk formed by the receptacle or some
part of it, or by a carpel. To distinguish its particular nature in any case,
the following terms are more or less employed :
THECAPHORE, for a stipe which belongs to a simple pistil itself (where
it is homologous with a petiole), and is no part of the receptacle, as in Coptis
or Goldthread.
GYNOPHORE, where the stipe is an internode of receptacle next below the
gynoecium, as the pod-stalk in some Cruciferae, Cleome, and Gynandropsis.
GONOPHORE, when it elevates both stamens and pistil, as it seemingly
does in the lower stipe of Gynandropsis, Fig. 409.
ANTHOPHORE, when the stipe is a developed internode between the
calyx and corolla, as in the Pink family, Fig. 408.
GYNOBASE is a term properly applied to a short and comparatively broad
portion of receptacle on which the gynoecium rests, as in Rue and Orange
(Fig. 414), Houndstongue, Sage, &c. This may extend up between the car-
pels and pass into, or the upper part become a
CARPOPHORE, a name properly applied to a portion of receptacle which
is prolonged between the carpels as a central axis, as in Geranium (Fig.
411) and many Umbelliferae, Fig. 412.
FIG. 408. Section of a flower of Silene Pennsylvanica, showing the stipe or
anthophore.
FIG. 409. Flower of Gynandropsis, with floral circles separated on the elongated
receptacle.
FORMS OF THE RECEPTACLE.
213
392. Instead of forming a stalk, the elongation may be continued
between the carpels in the form of a slender axis, as in Gera-
nium (Fig. 410, 411), and in the carpophore
of the fruit of Umbelliferse, Fig. 412. In
Geranium, this prolongation of receptacle
410
extends far above the ovaries as a beak, to which the styles arc
adnate for most of their length.
393. In Nelumbium (Fig. 413), the gynophore, or portion of
receptacle above the stamens, is enlarged into a singular broadly
top-shaped body, with a flat summit, in which the pistils (a dozen
or more isolated carpels) are separately immersed.
394. A Disk is a part of the receptacle, or a development of it,
enlarged under or around the pistil. When under it or around
its base and free from the calyx,
the disk is hypogynous, as in
Orange, Fig. 414. Here it is a
kind of g}'nobase. When adher-
ent to or lining the base of the
cah'x, it is perigynous, as in 415 416
Rhamnus (Fig. 415,416) and Cherry (Fig. 337) : when carried
by complete adnation up to the summit of the ovary, it is epigy-
nousi as in Cornus, in Umbelliferse, &c. Not rarely it divides
into lobes, as in Vitis (Fig. 379, 380), in Periwinkle and most
Apocj'naceous plants, and in Cruciferae. These are termed glands
of the disk, and indeed are commonl}* glandular or nectariferous.
FIG. 410. Gynoecium of Geranium maculatum. 411. The same with fruit mature,
the five ovaries or cells and the lower part of their styles separated and recurving away
from the prolongation of the axis or receptacle, to which they were at flowering-time
firmly attached.
FIG. 412. Mature fruit of Osmorrhiza. the two carpels splitting away below from
the filiform prolongation of the receptacle, or carpophore.
FIG 413. The top-shaped receptacle of Nelumbium, with the pistils, immersed in
hollows of its upper face.
FIG. 415. Flower of a Rhamnus or Buckthorn, and 416, section of the same, show-
ing a thickened perigynous disk.
214
THE FLOWER.
It is not possible by any direct demonstration to distinguish be-
tween such productions of the receptacle, which are classed as
belonging to the axis, and suppressed or undeveloped phyllous
organs, such as stamens, which glands of the disk may some-
times represent.
395. Hypanthium. Inspection of Fig. 415, 416, and 337, and
comparison with Fig. 339, will suggest an explanation differ-
ent from that which is generally
adopted. Instead of regarding the
calyx as beginning on a level with
the base of the ovary, and the cup
as lined, more or less thickly, by
an expansion of the receptacle (the
perigynous disk) ,
the calyx may be
understood to begin
where this and the
ovary become free
from each other.
Underthat view, the
receptacle, instead
of convex or protu-
berant, is here con-
cave, has grown up
419 around the ovaiy,
which, however, is free from the cup in the earlier cited figures,
but immersed in it in Fig. 339 and the like. A comparison with
a rose-hip, an apple, and a pear much strengthens this interpre-
tation, which is rather largely adopted at this day, at least
theoretically. It was perhaps first proposed by Link, who intro-
duced the appropriate name of HYPANTHUM. A hypanthium or
hypanthial receptacle is, as the name betokens, a flower-axis or
receptacle developed mainly under the calyx. The name is a
good one, in an}^ case ; and such structures as those of Calycan-
thus (Fig. 417-419), a rose, a pear (the lower part of which is
evidently an enlargement of peduncle), and of Cactus-flowers
(Fig. 317), although quite compatible with the theory of aclnation,
are more simply explained b}* it. 1
1 But, whether the cases are well distinguishable or not, it by no means
follows that the receptacle plays such a part in all instances of perigyny and
of inferior or partly inferior ovary. Such a view is attended by more diffi-
culties than the other. Unless the mediation of an invisible receptacle must
FIG. 417. Flowering branch of Calycanthus. 418. Vertical section of the urn -shaped
receptacle, the imbricated bracts or sepals on its surface cut away. 419. Mature
fructiferous receptacle entire, showing some scars from which the bracts have fallen.
ADAPTATIONS TO FERTILIZATION. 215
SECTION IV. CERTAIN ADAPTATIONS OF THE FLOWER TO THE
ACT OF FERTILIZATION.
1. IN GENERAL.
396. The introduction into morphological botany of the con-
siderations now to be mentioned should have dated from the
year 1793, in which Christian Conrad Sprengel published his
curious treatise on the structure of flowers in special reference
to insect aid in their fertilization. For this book, which was
wholly neglected and overlooked for more than sixty years, con-
tains along with some fanciful ideas the germs of the present
doctrine and many excellent illustrations of it. l The interest in
the doctrine now prevalent is witnessed by a copious special
literature, beginning with the publication, in 1862, of Darwin's
book on the fertilization of Orchids by the aid of insects. 2
be invoked whenever there is a junction of two dissimilar organs, the petals
and stamens of a Lythrum or a Cuphea are united with the calyx itself,
instead of calyx beginning at the top of a long and simple tube. And if
three or more of the floral whorls may be congenitally united, why not these
also with the remaining one ? Van Tieghem, in his Anatomie Comparee de
la Fleur, maintains wholly the old view, founding it upon anatomical struc-
ture and his ability to trace down to the base of the ovary the distinct
vascular bundles of the several involved organs.
1 C. C. Sprengel, Das entdeckte Geheimniss der Natur im Bau und der
Befruchtung der Blumen, Berlin, 1793. Even earlier, Koelreuter ( Vorlaufige
Nachricht, etc., 1761-1766) recognized the necessity of insect-aid to various
blossoms, and described some special contrivances for the purpose.
2 Charles Darwin, On the Various Contrivances by which British and
Foreign Orchids are fertilized by Insects, and on the Good Effects of Inter-
crossing, London, 1862. Ed. 2, 1877. This last contains a list of the papers
and books which bear upon the subject, published since 1862.
Other leading works and papers on the subject are, exclusive of the
other volumes and papers of Darwin, more or less referred to hereafter.
Treviranus, Ueber Dichogamie, &c., in Bot. Zeitung, xxi. 1863.
Hugo von Mohl, Einige Beobachtungen iiber dimorphe Blitthen, Bot.
Zeitung, xxi. 1863.
Delpino, Pensieri sulla Biologia Vegetale, &c., 1867. Kelazione sull'
Apparecchio della Fecondazione nelle Asclepiadie, &c., 1867. Ulteriore
Osservazioni sulla Dichogamia, &c., 1868-69, 1870, and later papers.
Axell, Om anordningarna for de Fanerogama Vaxternas Befruchtung,
Stockholm, 1869.
Hildebrand, Die Geschlechter-Vertheilung bei den Pflanzen, 1867, and
other papers.
Hermann MUller, Die Befruchtung der Blumen durch Insekten, 1873, and
papers in " Nature " and elsewhere.
"Flowers and their Unbidden Guests," an English translation of a work
by Professor Kerner, which describes arrangements in blossoms for exclud-
ing unwelcome guests, has not yet reached us. It introduces the new terms
Autogamy and Allogamy, defined on the following page ; the latter compre-
216 THE FLOWER.
397. The subject, here considered as a part of morphology,
must be fully treated, as regards acts and processes, under physi-
ology. Ever} r thing in the flower is in relation to fertilization
and fructification, directly or indirectly. This section is con-
cerned with those adaptations of structure by means of which
agents external to the blossom are brought into service for its
fertilization.
398. Linnaeus and his successors taught that the adjustments
in hermaphrodite flowers were such, on the whole, as to secure the
application of the pollen of its stamens to the stigma of its pistil
or pistils. The present view is, that this is doubtless strictly
secured in certain flowers of a moderate number of species, but
never in all the flowers of an}' such species ; that in ordinary
flowers, where it may commonly take place, it is not universal ;
that in the larger number of species there is something or other
in the floral structure which impedes or prevents it. Some
flowers are adapted for close fertilization ; some for cross fertili-
zation ; some for either. Here two terms need definition, viz. :
Close fertilization or Self-fertilization, or Autogamy, the applica-
tion and action of a flower's pollen upon its own pistil ;
Cross fertilization, or Allogamy, the action of the pollen of one
flower on the pistil of some other flower of the same species.
This may be near, as when between flowers borne in the same
cluster or on the same plant ; remote, when between flowers of
distinct plants of the same immediate parentage ; most remote,
when between different races of the same species. Any thing
beyond this is Irybridization, or crossing of species.
2. ADAPTATIONS FOR ALLOGAMY OR INTERCROSSING.
399. The doctrine now maintained appears to have been first
propounded by Sprengel in the statement that "Nature seems
to have wished that no flower should be fertilized by its own
pollen," a proposition which is not wholly tenable, for there
are blossoms specialty adapted to self-fertilization. It was re-
affirmed in our da}' by Darwin, in a similar adage, "Nature
abhors perpetual self-fertilization," a metaphorical expression
to which no effective exception has been taken. And the infer-
ence was drawn by him, that some important good to the species
must result from propagation through the union of distinct
individuals, and especially of individuals which have been dis-
tinct for several or many generations.
hencling Geitonoyamy, fertilization by pollen of other flowers of the same
plant, and Xenogamy, by pollen from a flower on another plant.
ADAPTATIONS FOR INTERCROSSING. 217
400. The actual proposition, simply stated, is that flowers are
habitually intercrossed, and that there are manifold structural
adaptations which secure or favor intercrossing, to such extent
as to justify the proposition. The prooi of the proposition is an
induction from a very great number of particular observations.
That intercrossing is beneficial is a rational inference from the
array of special adaptations for which no other sufficient reason
appears, or (to resume the metaphor) from the vast pains which
seem to have been taken to secure this end. This inference has
been to some extent confirmed by direct experiment. 1
401. Separation of the sexes is a direct adaptation to inter-
crossing, rendering it necessary between individuals in dioecious,
and largely favoring it in most monoecious and polygamous
flowers. Strictly close fertilization can occur in hermaphrodite
flowers only ; but it is in these that the most curious adaptations
for intercrossing are revealed.
402. The agencies to the one or the other of which most
flowers are structurally adapted in reference to intercrossing are
mainly two ; viz., the winds and animals, of these chiefly insects.
Delpino has accordingly classified flowers into Anemophilous and
Entomophiloiis ; literally wind-lovers and insect-lovers, but de-
noting wind-fertilized and insect-fertilized, according to the
agent by which pollen is transported. 2 There are hermaphrodite
and unisexual flowers of both classes, but most wind-fertilized
flowers are unisexual.
403. Wind-fertilizable or anemophilous flowers are mostly neu-
tral or dull in color, destitute of odor, and not nectariferous.
Their principal structural adaptations to this end, besides the
separation of the sexes in most of them, are the superabundance,
incoherency, dryness, and lightness of the pollen, rendering it
very transportable by wind and currents of air. The immense
abundance of pollen, its lightness, and its free and far diffusion
through the air in Pines, Firs, Taxodium, and other Coniferae,
are familiar. Their pollen fills the air of a forest during anthe-
sis ; and the u showers of sulphur," popularly so-called, the
3'ellow powder which after a transient shower accumulates as
a scum on the surface of water several or many miles from the
1 Darwin, The Effects of Cross and Self-Fertilization in the Vegetable
Kingdom, London, 1870. American Edition, New York, 1877.
2 Ornithophilons, i. e. bird-fertilized, flowers are to be ranked with entomo-
philous. The large blossoms of Trumpet Creeper (Tecoma radicans) and
of Trumpet Honeysuckle (Lonicera sempervirens), and others, are commonly
visited and probably fertilized by humming-birds as well as by moths ; and
other birds are known to play a similar part in equatorial regions.
218 THE FLOWER.
nearest source, testifies to these particulars. All amentaceous
trees (Willows excepted), Hemp, Hops, &c., are wind-fertilized ;
and, among perfect flowers, those of most Grasses, Sedges, and
Plantago. In the latter families especially, the anthers are pro-
truded or hung out in the air only when just ready to discharge
their pollen, and are at that moment suspended on suddenly
lengthened capillary drooping filaments, fluttering in the gentlest
breeze ; and the stigmas are either dissected into plumes, as in
most Grasses, or beset with copious hairs on which pollen is
caught. One physiological adaptation, very common in the fol-
lowing class, is not unknown among hermaphrodite wind-fertiliz-
able flowers, where it is important for securing intercrossing, viz.
Dichogamy. It is best seen in the common species of Plantago
or Plantain, and is described below. (408.)
404. Insect-fertilizable or entomophilous flowers are correlated
with showy coloration (including white, which is most show} 7 at
dusk), odor, or secretion of nectar, often by all three modes of
attraction to insects combined. Some insects, moreover, visit
flowers for their pollen, a highly nutritious article, and ordina-
rily produced in such abundance that much ma} r be spared.
The showiness of corolla or other floral envelopes is an attractive
adaptation to fertilization, enabling blossoms to be discerned at
a distance ; nor do we know that fragrance or other scent or
that nectar subserves any other uses to the flower than that of
alluring insects. Adaptations in the pollen of such blossoms
for transportation by insects are various. Common!}' the grains
are slightly moist or glutinous, or roughish, or studded with
projections, or strung with threads (as in CEnothera), so as not
to be readily dispersed in the air, but to have some slight
coherence as well as capability of adhering to the head, limbs,
or bodies of insects, especially to their rough surfaces ; and
in two families (Orchidaceae, Asclepiadaceae) the pollen is com-
bined in masses and with special adaptations for being trans-
ported en masse. (421.) With this the stigma is usually
correlated, by roughness, moisture, or glutinosity. 1
405. Adaptations of the flower itself in reference to insect
visitation are wonderfully various ; and most of these are found
upon investigation to favor, or often to necessitate, intercross-
ing. In difficious flowers, this is necessitated by the separation ;
in monoecious and polygamous flowers, of various kinds and
1 Thus nearly every Orchid genus but one has a persistently glutinous
stigma ; in the exceptional one, Cypripedium, it is moist and minutely rough-
ened, in correlation with the loosely granular or pultaceous pollen which it
is to receive.
ADAPTATIONS FOR INTERCROSSING. 219
degrees of separation, pollen is very commonly borne from
plant to plant ; in hermaphrodite flowers only are more special
arrangements needed to secure intercrossing or a certain measure
of it, and in these such arrangements abound.
406. Irregularity is one of the commonest modifications of the
flower (326, 337) : it is never conspicuous except in blossoms
visited by insects and generally fertilized by their aid ; and it
finds rational explanation on the score of utility in this regard. 1
407. Dichogamy, a term introduced by C. C. Sprengel, who
first noticed and described it, is one of the most usual and effect-
ual (rather physiological than morphological) adaptations for the
promotion of intercrossing between hermaphrodite flowers. It
means that such intercrossing is brought to pass by a difference
in the time of maturity of anthers and stigma ; this rendering
dichogamous blossoms practically the same as dioecious or mon-
oecious in respect to fertilization, while there is the economical
gain that all the flowers are fertile. According to whether the
anthers or the stigmas are precocious, dichogamous flowers are
Proterandrous (or Protandrous} , when the anthers mature and
discharge their pollen before the stigma of that blossom is recep-
tive of pollen ;
Proterogynous (or Protogynous} , when the stigmas are in
receptive condition before the anthers have matured their pollen.
Synanthesis^ 2 the maturing of the two sexes simultaneously or
nearly so, is however made to secure the same result through
special arrangements.
408. Proterogyny. The Plantains, such as Plantago major and
P. lanceolata, are familiar instances of this in a wind-fertilized
genus with hermaphrodite flowers. The anthesis proceeds from
base to apex of the spike in regular order, and rather slowly.
While the anthers are still in the unopened corolla and on short
filaments, the long and slender hairy stigma projects from the tip
and is receiving pollen blown to it from neighboring plants or
1 This did not escape the attention of Sprengel in the last century, and
along with it the fact that strictly terminal and also vertical flowers, whether
erect or suspended, are seldom irregular, while comparatively horizontal or
obliquely set flowers more commonly are so. The irregularity is in refer-
ence to a landing place for the visiting insect, or also to storage of or accessi-
bility to nectar, &c.
Darwin (Forms of Flowers, 147) remarks that he does not know of a
single instance of an irregular flower which is wind-fertilized.
2 Synacmy is the term proposed by A. W. Bennett, in Journal of Botany,
viii. (1870), 316, with its opposite, Ifeteracmy, for proterandry and proterogyny.
The latter names, in their shorter form (protandry and protogyny), appear to
have originated with Hildebrand, 1867.
220 THE FLOWER.
spikes : a day or two afterwards, the corolla opens, the filaments
greatly lengthen, and the four anthers now pendent from them
give their light pollen to the wind ; but the stigmas of that flower
and of all below it on that spike are withered or past receiving
pollen. Among Grasses, Anthoxanthiun is in the same case.
The arrangement is somewhat similar to the Plantain in Arnor-
plia, which is fertilized by insects, the simple stigma projecting
beyond the corolla in bud, while the anthers are still immature
and enclosed. Scrophularia is a good instance of proterogyny
in flowers fertilized by bees. The flower is irregular (Fig.
420-422) , and is approached from the front, the spreading lower
lobe being the landing place. Fig. 420 represents a freshly
opened blossom : and Fig. 421, a section of it. Only the style
tipped with the stigma is in view, leaning over the landing place ;
the still closed anthers are ensconced below. The next day or
a little later all is as in Fig. 422. The style, now flabby, has
fallen upon the front lobe, its stigma dry and no longer receptive :
the now-opening anthers are brought upward and forward to the
position which the stigma occupied before. A honey-bee, taking
nectar from the bottom of the corolla, will be dusted with pollen
from the later flower, and on passing to one in the earlier state
will deposit some of it on its fresh stigma. Self-fertilization
here can hardly ever take place, and only through some disturb-
ance of the natural course.
409. Proterandry. The process is the reverse, and is at-
tended with much more extended movements in Clerodendron
Thompson!, a Verbenaceous tropical African climber now com-
mon in conservatories. The adaptations in this flower" (which
we indicated long ago) are exquisite. The crimson corolla
and bright white calyx in combination are very conspicuous.
The long filiform filaments and style, upwardly enrolled in the
FIG. 420, 421. Early opened flower of Scrophularia nodosa, and a longitudinal
section. 422. Flower a day or two later.
ADAPTATIONS FOK INTERCROSSING. 221
bud, straighten and project when the corolla opens : the stamens
remain straight, but the style proceeds to curve downward and
backward, as in Fig. 423. The anthers are now discharging
pollen : the stigmas are immature and closed. Fig. 424 repre-
sents the flower on the second day, the anthers effete, and the
filaments recurved and rolled up spirally ; while the style has
taken the position of the filaments, and the two stigmas now
separated and receptive are in the very position of the anthers the
previous day. The entrance by which the proboscis of a butterfly
may reach the nectar at bottom is at the upper side of the orifice.
The flower cannot self- fertilize. A good-sized insect ftying from
blossom to blossom, and plant to plant, must transport pollen
from the one to the stigma of the other.
410. Proterandry abounds among common flowers. It is
conspicuous in Gentians and in nearly all that family. But,
while in Gentians the short style is immovable and erect, in
Sabbatia it is thrown strongly to one side, out of the way of and
far below the stamens, the branches closed and often twisted, so
that the stigma is quite inaccessible until the stamens have shed
their pollen : then the style becomes erect, untwists, its two flat
branches separate, and expose the stigmatic surface of their inner
face in the place which the anthers occupied. In Sabbatia
angularis, Lester F. Ward 1 observed that the anthers of freshly
1 In Meehan's Gardeners' Monthly, September, 1878, 278.
FIG. 423. Flower of Clerodendron Thompsoniae, first day ; 424, second day.
2:22
THE FLOWER.
opened blossoms are all thrown to one side almost as strongly
as the st}*le is thrown in the opposite direction. One of our
common Fireweeds, Epilobium angustifoliimi or E. spicatum, as
it is variously called, which is common all round the northern
hemisphere, is similar to Sabbatia in behavior. In the freshly
opened flower, while the anthers are in good condition and are
giving their pollen to bees, the still immature style is strongly
curved downward and backward, as in Fig. 425. Two or three
days later, when the pollen is mostly shed, the style straightens,
lengthens to its full dimensions, and spreads its four stigmas over
the line of the axis of the blossom (Fig. 426), in the very
position to be pollinated by a bee coming from an earlier flower.
411. In the following instances of proterandry, the st} T le is
made the instrument of distributing the pollen which it is not
itself to use. The
anthers of a Cam-
panula discharge all
their pollen in the
unopened bud, and it
is nearly all deposited
on the style which
they surround, the
upper part of which
is clothed with a coat
of hairs for holding
427 428 the pollen. (Fig. 427.)
In the open flower, the stamens are found to be empty and withered,
as in Fig. 428. These flowers are visited by bees and other insects
for the pollen. While this is going on, and while the pollen is
fresh and plentiful, no stigma is apparent. Later, the top of the
st}*le opens into three (in some species five) short and spreading
branches, the inner faces of which are the stigmas. Although
FIG. 425, 426. Flowers of Epilobium angustifolium or spicatum; in the first,
freshly expanded; in the second, a few days older.
FIG. 427. Vertical section of an unopened flower of Campanula rapunculoides :
the broad white lines are sections of two anthers. 428. Same of an older flower.
ADAPTATIONS FOR INTERCROSSING.
223
so close at hand, little if any of the pollen of that flower can
reach the stigmas. These actually get fertilized by pollen
brought by bees, which come loaded with it from
other flowers and other plants. Symphyaadra differs
from a true Campanula chiefly in the continued
cohesion of the five anthers into a tube around the
style. (Fig. 429, 430.) The pollen is discharged on
and held by the hairy upper portion of the style.
Soon after, the corolla expands, the lower
part of the style lengthens, and carries
the pollen-loaded part out of and above
the anther- tube, as in Fig. 430 ; lastly, the
three connivent tips of the style diverge
and expose the stigmas to pollen mainly
brought by bees from other flowers. B}' a
slight further modification in Lobelia and
in Composite, pollen is pushed out of the
anther-tube b} r the tip of the style as it
lengthens, or by the very back of the two
stigmas, the faces of which, afterwards
exposed, are not to receive this, but other 429 *"
pollen, though it may at times receive some of its own. The
arrangement in Composite is here illustrated from Leptosyne
maritima (Fig. 431-435), a showy plant of Southern California,
now not veiy rare in cultivation. The large flowers around the
FIG. 420. Stamens and pistil of a young, and 430, same from an old flower of
Symnliyandra pendula
FIG. 431. Head of flowers of Leptosyne man" Una. of the natural size.
224
THE FLOWER.
margin (ray-flowers, with ligulate corolla) , one of which is sepa-
rately shown in Fig. 432, are pistillats only : the enlarged and
extended open part of the
corolla (bright yellow in
color) serves for attrac-
tion, the circle of rays
gives the appearance as
of a single large flower.
The flowers of the disk
or whole central part are
hermaphrodite, and with
narrow tubular corollas,
from the orifice of which
projects the greater part
of the tube of five eo-
alescent anthers. The pollen is early discharged into the interior
of this tube. The style, with somewhat enlarged and brush-like
tip, at first reaches only to the
bottom of the anther-tube : it
slowly lengthens, pushes the
pollen before it out of the tube
(Fig. 433) and into the way of
insects of various kind, which,
travelling over the surface, con-
vey it to older flowers of the same
head and of other plants. The style, elongating yet more, raises
some of the pollen still higher (as in Fig. 434) ; and at length its
two branches separate and diverge (Fig. 435), exposing to other
pollen the stigmatic receptive surface which until now was un-
approachable.
412. In Parnassia, which has sessile stigmas, their receptive
surface is actually not formed until the anthers become effete ;
FIG. 432 A ligulate female flower of the same, and a, central hermaphrodite flower.
433. Upper part of the latter, more enlarged, the tube of anthers projecting from the
corolla, and the pollen projecting from apex of the anther-tube, being pushed up by the
lengthening of the style beneath. 434. This style now projecting, and some pollen still
resting on its tip. 435. Tip of same style (more advanced and magnified); the two
branches spreading, still carrying some pollen on the apex of each arm or branch, and
by the divergence now exposing the stigmatic inner faces.
ADAPTATIONS FOR INTERCROSSING.
225
and, as the plants or stems are single-flowered, the}' are function-
all)' dioecious while structurally hermaphrodite.
413. The adaptations for hermaphrodite intercrossing with
synanthesis (407) , i. e. where there is no essential difference of
time in the maturing of anthers and stigma, are manifold.
The}' may be classed into those without and those with dimor-
phism of stamens and pistils, or, in other words, those with
Homogenous and those with Heterogonous flowers. 1
414. The cases without dimorphism are the most various,
certain families having special types ; and are of all degrees,
from those that require intercrossing to those that merely favor
or permit it. For the present purpose, having only morphology
in view, it suffices to bring to view two or three cases or types of
415. Particular Adaptations in hermaphrodite blossoms, not
involving either dichogamy or dimorphism. These are exceed-
ingly various ; but they may be distinguished into two general
kinds, namely: 1, where loose and
powdery pollen is transported from
blossom to blossom in separate grains,
and 2, where pollen-masses or the
whole contents of anthers are bodily
so transported.
416. Papilionaceous flowers (such
as pea-blossoms, 338) having ten
stamens enclosed with a single pis-
til in the keel
of the corolla,
their anthers in
close proximity
4T8
to the stigma
were naturally
supposed to be
self-fertilizing ; and so they sometimes are, jet with marked
adaptations for intercrossing. None are less so than those of
1 Terms proposed in Amer. Jour. Sci. ser. 3, xiii. 82, and in Amer.
Naturalist, January, 1877. Dimorphism in flowers may affect the perianth
only, and not the yovf) or essential organs ; or there may be two kinds of
flowers as respects these also, but with no reciprocal relations, as in cleisto-
yamous dimorphism (534) ; or of two kinds essentially alike except in stamens
and pistil, and these reciprocally adapted to each other, which is heterogonous
dimorphism, or, when of three kinds, trimorphism.
FIG. 436. Flower of Wistaria Sinensis natural size. 437. Same enlarged, with
standard, wings, and half the keel removed. 438. Same with the keel depressed, as it
is when a bee alights on this its usual landing place, the cluster of anthers and stigma
thus brought up against the bee's abdomen. 439. Style and stigma, with part of the
ovary, more magnified, a fringe of fine bristles around the stigma.
15
226
THE FLOWER.
442
Wistaria (Fig. 436-439), in which the light fringe of stiff hairs
around the stigma (shown in Fig. 430) would not prevent pollen
of surrounding anthers from falling upon it. Yet when a bee
alights upon the keel,
with head toward the
base of the flower, and
proboscis is inserted
for nectar between the
foot of the standard
and the keel, the latter
is depressed by the
weight, so that the ab-
domen of the insect is
brought against the ten
anthers and the stig-
ma, becoming thereby
smeared with pollen,
some of which wh^n
other blossoms are vis-
ited cannot fail to be applied to their stigmas. The very similar
flower of Locust (Robinia), like that of the Pea, adds an adapta-
tion in favor of intercrossing. The style for some length below the
stigma is covered with a short beard of hairs, as is
seen in Fig. 442. The anthers open earl}' and dis-
charge their pollen, which mainly lodges on this
beard (Fig. 443), in a manner which may thus far
be likened to the case of Campanula. (411.) The
wings and the keel are yoked together, and are
together depressed by the weight of an alighting
44-5 bee. This does not bring out the anthers as in
Wistaria, but these remain until effete within the sac, while the
stigma and the pollen-laden part of the style (Fig. 441) are
projected against the bee's abdomen, which, b}- the oblique
movement, is first touched by the stigma and next brushed over
with pollen by the st3'le below. So that, in visiting a succession
of blossoms, some pollen of one flower is transferred to the body
of the bee, and thence to the stigma of the next flower, which
flower immediately gives to the same spot some of its pollen, to
be transferred to the next flower's stigma, and so on.
417. Two special modifications of the papilionaceous t}*pe
FIG. 440. Flower of Robinia hispida. the standard and wings removed. 441. Same,
as depressed by the weight of a bee, causing the stigma and pollen-laden tip of the style
to protrude. 442. Enlarged section of same in the bud, leaving one keel-petal, half the
stamens, and the pistil in view. 443. Style and stigrna at a later period, the beard
loaded with pollen; more magnified.
ADAPTATIONS FOE IXTEllUKOSSiNG. '2'21
need particular mention. One of them, the Bean-blossom, is
well known to botanists ; the other not so. The peculiarity in
the common Bean, Phaseolus vulgaris, and its nearest relatives,
is that the keel, enclosing the stamens and pistil, is prolonged
into a narrow snout which is spirally coiled (as in Fig. 444-446) ;
that the stigma is oblique on the tip of the style, and the beard
on the style is mainly on the same side that the stigma is : the
wing-petals stand forward and turn downward, forming a con-
venient landing place for bees. As in the Locust-blossom, the
anthers early discharge their pollen, much of which adheres
lightly to the beard of the style. In the untouched flower, all
from first to last is concealed in the coiled keel. Press down the
wing-petals, and first the stigma and then the pollen-laden tip
of the style projects from the orifice : remove the pressure, and
the}' withdraw within. When this pressure is made by a bee,
resting on the wing-petals while searching for nectar within the
base of the blossom between the keel and the standard, the same
movement occurs : the stigma first, and then the pollen on the
style, strikes against a certain portion of the front or side of
the bee's body, and the repetition of this operation causes the
fertilization of each blossom by other than its own pollen. A
slighter pressure or lighter movement of the wing-petals suffices
FIG. 444. Flower of Garden Bean. Phaseolns vulgaris. 445 Same with wing-
petals pressed down and tip of style projecting from the orifice of the keel. 446. Same
as 444 enlarged, and standard and wings removed 447. Upper part of keel, in the
condition of 445, enlarged, showing plainly the projecting style. 448 Section of the
keel, enlarged, showing the style within before the anthers open: stamens for sake of
clearness not delineated. 449. Pistil detached from an older flower ; the brush loaded
with pollen.
228
THE FLOWER.
to jostle some of the pollen down upon its own stigma, so that
self-fertilization is not uncommon.
418. Apios tuberosa, a near relative of Phaseolus, exhibits a
different and equally curious modification of the same parts.
^ ie wm S~P eta l s f r landing place
are similar: the standard is pro-
portionally large, firm in texture,
and shell-shaped or concave, with
a small boss at the tip as seen from
behind, or a shallow sac as seen
450 ^ from the front : the keel is narrow
and sickle-shaped ; it arches across the front of the flower, and
the blunt apex rests in the notch or shallow sac of the tip of Hie
standard. (Fig. 450, 452, 453.) So it remains if untouched until
the blossom withers : no self-fertilization has ever been observed,
and none ordinarily occurs. The anthers
are assembled close around the stigma,
but a little short of it (Fig. 452) ; the
pollen is not early nor copiously shed in
the enclosure : the small terminal stigma
is at first covered with a pulp3 T secretion,
which at length collects into a soft ring
around its base over or through which no
pollen passes. But when the keel is liber-
ated by lifting from underneath, it curves
promptly into the shape shown in Fig.
PIG 450. Flower of Apios tuberosa, unvisited. 451. Same after visitation, the
keel dislodged from the retaining notch, and more incurved; the tip of the style pro-
truded and thrust forward, followed by the anthers.
FIG. 452. Enlarged vertical section of flower-bud of Apios tuberosa. 453 A flower
wjth half the standard cut away, to show the blunt apex of the keel resting in the
notch. 454. Diagram of flower, with half of the standard cut away, to show what takes
place when the apex of the keel is liberated. The figures (also those from 423 to the
present), and t,h> first account of the adaptations of Apios, were published in the Amer-
ican Agriculturist in 1876.
ADAPTATIONS FOR INTERCROSSING. 229
451, or better in Fig. 454, where the dotted lines indicate its
original position ; and first the end of the style, tipped with
its stigma, is pushed forward, and then the anthers come into
view. The flowers are visited by humble-bees, and sometimes
by hone} T -bees. In searching for nectar at the base of the flower,
they probably push forward into the space under the arching
keel, and by slightly elevating dislodge its apex ; when first the
stigma and then the anthers are brought against some portion
of the insect's body, and against the same portion in succeeding
blossoms, thus effecting cross-fertilization. This rationally ex-
plains a remarkable adaptation, which seems to be not otherwise
intelligible.
419. Special Adaptations. Two of these, each peculiar to
the genus, may here be referred to. In Kalmia-blossoms (Fig.
455 456 457
455-458), the anthers discharge the pollen through a small
orifice at the apex of each cell, in this respect agreeing with
Rhododendrons and their other relatives ; but
none of them utilize this family peculiarity in the
manner of Kalmia. In the flower-bud, each of the
ten anthers is lodged in a small cavity or pocket
(externally a boss) of the corolla, in a way analo-
gous to that in which the keel of Apios is lodged in
the tip of the standard (418) : the expansion of the
border of the corolla in anthesis curves the fila-
ments outward and backward ; and when the bowed 4.53
stamens are liberated by rough jostling the}' fly up elastically,
and the pollen is projected from the two orifices. Some pollen
may possibly be thrown upon the single small stigrna at the
tip of the style, which rises much above the stamens. But the
anthers are not dislodged when undisturbed, at least until
after the elasticity of the filaments is lost : they are dislodged by
humble-bees, which circle on the wing over the blossom, the
FIG. 455 Vertical section of a flower-bud of Kalmia latifolia, showing the anthers
lodged in the pockets of the corolla. 456. Expanded flower, with bowed stamens.
"\T. Vertical section of the same. 458. A stamen, enlarged.
230
THE FLOWER.
under side of the abdomen frequently touching the stigma, while
the proboscis is searching round the bottom of the flower, liberat-
ing the stamens in the process, which one by one project their
pollen upon the under side of the insect's bodj'. In the passage
from flow r er to flower, pollen is thus conveyed from the anthers
of one to the stigma of another.
420. Iris has three stamens, one before each sepal or outer
lobe of the perianth, and behind each petal-like lobe of the style
(Fig. 459) : the stigma, a
shelf-like plate of each lobe,
is just above the anther ;
but, as the anther faces
outward and the stigma is
higher and faces inward, no
pollen can find its way from
the one to the other. But the
adaptation of parts is admir-
able for conveyance by bees,
which, standing upon the
only landing place, the re-
curved sepal, thrust the head
down below the anther, and
in raising it cany off pollen,
to be afterwards lodged
upon the stigmas of other
flowers which they visit.
421. Transportation of Pollinia, or of all the pollen in a
mass, is effected in most of the species of two large orders,
not otherwise allied, the Asclepiadaceae and the Orchidacese.
While in the Iris family the number of stamens is reduced from
six to three, in all the Orchis family, except Cypripedium, the
stamens are further reduced to a single one ; but the pollen is
peculiarly economized. That of Arethusa is in four loose and
soft pellets, in an inverted casque-shaped case, hinged at the
back, resting on a shelf, the lower face of which is glutinous
stigma, over the front edge of which the casque-shaped anther
slightly projects ; and this anther is raised by the head of a bee
when escaping out of the gorge of the flower. The loose pellets
of pollen are caught upon the bee's head, to the rough sur-
face of which they are liable 1 to adhere lightly and so to be carried
to the flower of another individual, there left upon its glutinous
FIG. 459. Flower of Iris puniila, with front portion and half of one petaloid style-
lobe and stigma cut away. The section of the stigma is seen edgewise: the rough
upper surface only is st
ADAPTATIONS FOR INTERCROSSING.
231
stigma by the same upward movement which immediately after-
ward raises the anther-lid and carries away its pollen, to be
transferred to a third blossom, and so on.
422. But it is in Orchis
and in the commoner re-
presentatives of Orchis in
North America (viz. Ha-
benaria, &c.) that the
most exquisite adapta-
tions are found, and the
greatest econom}^ se-
cured ; paralleled, how-
ever, by most of the very
numerous and various epi-
phytic and by various ter-
restrial Orchids of warmer
regions. A single illus-
tration ma}' here suffice ;
and Darwin's volume on
the Fertilization of Or-
chids (396, note), with
its references to the
copious literature of the
subject, may be studied
for full particulars and
their bearings. The flower
is trimerous, and the peri-
anth adnate to the ovary,
therefore apparent!}' de-
veloped upon its sum-
mit. The three external
parts of the perianth,
which in Habenaria orbi-
culata (Fig. 460) are
much the broader, are the
sepals: the three alternate
and internal, the petals :
the base of the long and narrow petal which is turned downward
is hollowed out and extended below into a long tube, closed at
bottom, open at top (the spur or nectary), in which nectar is
FIG 400. Flower of Habenaria or Platantheraorbiculata. enlarged. 461. Combined
stamen and stigma, more enlarged. 462. One of the two pollen-masses (poflinia), \vith
its stalk and glutinous disk or gland. 462. Lower part of this stalk and its disk, more
magnified.
232
THE FLOWER.
copiously secreted and contained. The central part of the
blossom, bej'ond the orifice of the nectary (shown separately in
Fig. 461), consists of one anther and a stigma, fused together
(the clinandrium) : the marginal portions, opening by a long
chink, are the two cells of the anther, approximate at their
broader portion above, widely divergent below : most of the
lower part of the space between is excessively glutinous, and is
the stigma. The grains of pollen are united by means of short
threads of very elastic tissue into small masses, and these into
larger, and at length into pellets, having stalks of the same
elastic tissue, b}^ which they are
all attached to a firmer central
stalk, or caudicle. (Fig. 4G3-465.)
To the lower end of this caudicle
(directly to the end of it in our
Habenariae and Orchises gener-
ally, in this instance to the inner
side of the end, with a thick inter-
mediate base intervening), is at-
tached a button-shaped disk, the
face of which is exposed, and is
on a line with the surface of the
anther ; so that these two disks
look toward each other across the
broad intervening stlgmatic space,
as seen in Fig. 4G1. The exposed
face of the disk being covered with
a durable layer of very viscid mat-
ter, the body itself is sometimes termed a gland, and not improperly.
The viscidit}' is nearl}* of the same nature as that of the interven-
ing stigma, of which the glands are generally supposed to be
detached portions. If so, then a portion of the stigma is cut off
from the rest and specialized to the purpose of conveyance of the
pollen. When a finger's end or any smaller body is touched to
these disks, the}' adhere so firmly that the attached pollinia or
pollen-masses are dragged out of the cell and carried away en-
tire. Some of these pollen-masses have been found attached by
the disk to the eyes of a large moth. When a moth of the size
of head and length of proboscis of Sphynx drupiferarum visits a
spike of these flowers, and presses its head into the centre of the
FIG. 463. A more magnified polle:)-mass of Platanthora orbieulata. with its stalk
and gland. 464 Five of the separate porti-nis or pollen-packets, \vith some of the
elastic threads of tissue eonneetina: them 465. A portion more highly magnified, with
some of the pollen-grains in fours detached.
ADAPTATIONS FOR INTERCROSSING.
23;]
flower so that its proboscis may reach and drain the bottom of
the nectariferous tube, a pollen-mass will usually be affixed to
each eye : on withdrawal,
these will stand as in Fig.
466. Within a minute they
will be turned downward
(Fig. 466), not by their
weight, but by a contraction
in diying of one side of the
thick piece which connects
the disk with the stalk.
When a moth in this con-
dition passes from the last
open flower of one spike to
that of another plant, and
thrusts its proboscis down a
nectar}', the transported pol-
len-masses will be brought
in contact with the large
glutinous stigma : on with-
drawal, either some of the
small pellets of pollen will be
left adherent to the stigma,
the connecting elastic threads
giving way ; or else a whole
pollen-mass will be so left,
its adhesion to the glutinous 41J a
stigma being greater than that of the disk to the moth's eye.
The former is a common and a more economical proceeding, as
then a succession of flowers are abundantly fertilized by one
or two pollen-masses. In either case, new pollen-masses arc
carried off from fresh flowers and applied to the fertilization
of other blossoms on the same and eventually on those of differ-
ent individuals. Cases like this, and hundreds more, all equally
remarkable, serve to show how sedulous, sure, and economical are
the adaptations and processes of Nature for the intercrossing of
hermaphrodite flowers.
422". An arrangement analogous to that of Orchids, and
similarly subservient to cross-fertilization, characterizes the
otherwise widely unlike Asclepias family. In Asclepias (Milk-
weed) there are five stamens surrounding a large stigmatic
PIG. 466 Front part of Sphynx drupiferarum. bearing a pollen-mass of Platan-
tliera orhionlata affixed to each eye, in the early positi-m. 466 <*. Front view of the head.
later, showing the pollen-masses deflexed.
234 THE FLOWER.
body, and alternating with these five two-cleft glands, the ver-
tical chink or groove of which is glutinous. To each gland is
firmty attached, by a cauclicle or stalk, a pollen-mass of an ad-
jacent anther. (Fig. 522.) A slight force upraising the gland
detaches it from the stigma and drags the pair of suspended
pollen-masses out of their cells. Insects visiting the blossoms
commonly dislodge them, the gland adhering to their legs or
tongues when these happen to be drawn through the adhesive
chink, and convey them from one flower to another. Without
such aid the flowers of Asclepias rarely set seed. 1
423. Dimorphism, t. e. the case of two kinds of blossoms, both
hermaphrodite, on the same species, is another adaptation to
intercrossing. Not all dimorphism, however, for in clristogamous
dimorphism (434) the intent to self-fertilize is evident. There
may also be dimorphism as to the perianth, not particularly
affecting fertilization. One kind, however, and the commonest,
is a special adaptation to intercrossing, viz. :
424. Heterogonous Dimorphism. (413, note.) This term is
applied to the case in which a species produces two kinds of
hermaphrodite flowers, occupying different individuals, the flowers
essentially similar except in the andrcecium and gynoecium, but
these reciprocally different in length or height, and the adapta-
tions such that, by the agency of insects, the pollen from the
stamens of the one sort reciprocally fertilizes the stigma of the
other. 2 This dimorphism has been detected in about forty genera
belonging to fourteen or fifteen natural orders, widely scattered
through the vegetable kingdom ; but there are far more examples
among the Rubiaceae than in any other order. Sometimes all
the species of a genus are heterogonous, as in Houstonia, and
1 The reported sensitiveness of the gland, referred to in the first issue of
this volume (1879), was founded upon misinterpreted observations.
2 This peculiar arrangement has been long known in a few plants, such
as Primula veris, P. grandiflora, and Houstonia. In Torrey and Gray's
Flora of North America, ii. 38, 39 (1843), these flowers are said to be dioecio-
dimorphous, not denoting that they are at all unisexual, but that the two
forms occupy different individuals. Their meaning was detected by C.
Darwin, and made known in his paper "On the Two Forms or Dimorphic
Condition in the Species of Primula, and on their Remarkable Sexual Kela-
tions," published in the Journal of the Linnean Society, vi. (1862), 77 : rcpub-
lished, in 1877, as the leading chapter of his volume entitled "The Different
Forms of Flowers on Plants of the Same Species." Mr. Darwin had termed
these flowers simply Diinor/tJn'c ; but in this volume he adopted Hilde-
brand's name of Heterostyled for this kind of blossom. The difference,
however, affects the androecium, and even the pollen, as well as the style ;
wherefore we proposed for it the name of Heterogonous or Heteroyone dimor-
phism, as mentioned in a former note, 413.
ADAPTATIONS FOR INTERCROSSING. 235
Cinchona, sometimes only a part of them, as in Primula and
Linum. In Hottonia, a Primulaceous genus of two species, the
European one has heterogonous dimorphism l for cross-fertiliza-
tion : the American one has homogenous show}' flowers with
only the general chance for intercrossing, and earlier flowers
which are cleistogamous for self-fertilization.
425. The nature of heterogone dimorphism may be well under-
stood from a single example. The most familiar one is that of
Iloustonia ; but, in larger blossoms, Gelsemium is a fine illus-
tration in the Southern United States, and Mitchella (Fig. 467)
mostly in the Northern. Raised from the seed, the individuals
are about equally divided between the two forms : namely, one
f jrai with bug style and short or low-inserted stamens ; the
other with short style and long or high-inserted stamens. The
stigmas in one rise to about the same height as the stamens in
the other, both in the tall or exserted organs and in their low
and included counterparts, as is shown in Fig. 468, answering
to the left hand and Fig. 469 to the right hand flowers of Fig. 467.
A bee or other insect with proboscis of about the length of the
corolla-tube, visiting the blossoms of Mitchella, will brush the
same part of its body against the high anthers of the long-
stamcned and the high stigmas of the long-styled forms ; and
1 C. C. Sprengel, as Darwin mentions, had noticed this, before 1793. He,
" with his usual sagacity, adds that he does not believe the existence of
the two forms to be accidental, though he cannot explain their purpose."
Darwin, Forms of Flowers, 51.
Some heterogonous Primulas are said to produce homogenous varieties in
cultivation. In Primula, and in other genera, there arc species Avhich seem
as if of one sort only, no reciprocal sort being known, as if one form had
become self-fertile and the other had disappeared.
FIG. 467. Partridge Berry, Mitchella repens. in the two forms, viz. long-stamenetl
and sliort-stylecl, aivl short-stamenel and lon~-stylc 1.
230
THE FLOWER.
the same part of the proboscis against the low anthers of the
short-stamened and the low stigmas of the short-styled form.
426. Moreover, Dar-
u in has ascertained by
microscopical examina-
tion that the pollen of
the two differs in size or
shape, and by experi-
ment that it is less
active upon its own
stigma than upon the
other ; indeed, that in
man}' cases (as in some
species of Linum) it is
quite inactive or impo-
tent not only upon its
own stigma but upon its
own-form stigma, while
it is prepotent on the other, and this reciprocally of the two
forms. 1 Here, then, are flowers structurally hermaphrodite, but
functionally as if dioecious, securing all the advantages of the
latter, along with the economical advantage that both sorts of
individual and every blossom may bear seed. With dicecism
only about half the plants could be fruitful.
427. Heterogonous Trimorphism. A threefold heterogonism
is known in certain species of a few genera ; and this complica-
tion may have certain conceivable advantages over dimorphism.
Where seedling dimorphous individuals are few and far between
(those multiplying from root would all be alike), there would
be an even chance that any two near each other were of the
same form and therefore sterile or imperfectly fertile. But if
the organization were of three forms, any two of which inter-
crossed with perfect fertility, the chances (as Darwin remarks)
are two to one that any two plants were of different forms, and
therefore by fertilizing each other completely fruitful.
428. The earliest known instance of three forms as to recip-
rocal relative length of stamens and pistil is that of Ly thrum
1 Impotence of own pollen, either absolute or relative, occurs no less in
certain flowers which are not dimorphous, as in Corydalis, some species of
Passiflora, &c. On the contrary, many dimorphous flowers are in a certain
degree self-fertile, especially in the long-stamened and short-styled form.
These subjects are physiological, and belong to another volume.
FIG. 468. Long-styled flower of Fig. 467. laid open. 469. Long-staraened flower of
the same laid open. Both equally enlarged.
ADAPTATIONS FOR INTERCROSSING.
237
Salicaria. This was indicated by Vauclier in 1841, more par-
ticularly described by Wirtgen in 1848, but was interpreted by
Darwin, and the more recondite differences brought to notice, in
1864. 1 " The three forms may be conveniently called, from the
unequal length of their pistils, the long-styled, mid-styled, and
short-styled. The stamens also are of unequal lengths, and
these may be called the longest, mid-length, and shortest."
The pollen of the different classes of stamens is of two sorts as
to color, and of three as to size, the largest grains from the
largest stamens. " The pistil in each form differs from that in
either of the other forms,
and in each there are two
sets of stamens, different
in appearance and func-
tion. But one set of
stamens in each form
corresponds with a set
in one of the other two
forms. Altogether, this
one species includes three
females or female organs,
and three sets of male
organs, all as distinct
from one another as if
they belonged to different
species ; and, if smaller
functional differences are
considered, there are five
distinct sets of males. Two of the three hermaphrodites must
coexist, and pollen must be carried by insects reciprocall}'
from one to the other, in order that either of the two should be
fully fertile ; but, unless all three forms coexist, two sets of
stamens will be wasted, and the organization of the species as
a whole will be incomplete. On the other hand, when all three
hermaphrodites coexist, and pollen is carried from one to the
other, the scheme is perfect : there is no waste of pollen and no
1 In an article On the Sexual Relations of the Three Forms of Lythrum
Salicaria, in Jour. Linn. Soc. viii. 169. Also on the Character and Hybrid-
like Nature of the Offspring of the Illegitimate Unions of Dimorphic and
Trimorphic Plants. Ihid. x. 393, 1868. Reproduced and extended in his
volume entitled " Forms of Flowers," 1877.
FIG. 470. Diagram of the flowers of the three forms of Ly thrum Salicaria, in their
natural position, with the petals and calyx removed on the near side. The dotted
lines with the arrows show the directions in which pollen must be carried to each
stigma to ensure full fertility. (From Darwin.)
470
238 THE FLOWER.
false coadaptation." The whole arrangement is clisplaj'ed in
the annexed diagram (Fig. 470), and in the following account
of the operation. 1 " In a state of nature, the flowers are inces-
santly visited for their nectar by hive and other bees, various
Diptera, and Lepidoptera. The nectar is secreted all round the
base of the ova Hum ; but a passage is formed along the upper
and inner side of the flower by the lateral deflection (not repre-
sented in the diagram) of the basal portions of the filaments ;
so that insects invariably alight on the projecting stamens and
pistil and insert their proboscides along the upper and inner
margin of the corolla. We can now see why the ends of the
stamens with their anthers and the end of the pistil with the
stigma are a little upturned, so that they may be brushed by
the lower hairy surfaces of the insects' bodies. The shortest
stamens, which lie enclosed within the calyx of the long- and
mid-styled forms can be touched only by the proboscis and narrow
chin of a bee : hence they have their ends more upturned, and
they are graduated in length, so as to fall into a narrow file,
sure to be raked by the thin intruding proboscis. The anthers
of the longer stamens stand laterally farther apart and are more
nearly on the same level, for they have to brush against the
whole breadth of the insect's body. . . Now I have found no
exception to the rule that, when the stamens and pistil are bent,
they bend to that side of the flower which secretes nectar. . . .
When nectar is secreted on all sides, the} r bend to that side
where the structure of the flower allows the easiest access to it,
as in Lythrum. ... In each of the three forms, two sets of sta-
mens correspond in length with the pistil in the other two forms.
When bees suck the flowers, the anthers of the longest stamens,
bearing the green pollen, are rubbed against the abdomen and
the inner sides of the hind legs, as is likewise the stigma of the
long-styled form. The anthers of the mid-length stamens and
the stigma of the mid-styled form are rubbed against the under
side of the thorax and between the front pair of legs. And,
lastly, the anthers of the shortest stamens and the stigma of the
cliort-styled form are rubbed against the proboscis and chin ; for
the bees in sucking the flowers insert onh r the front part of their
heads into the flower. On catching bees, I observed much green
pollen on the inner sides of the hind legs and on the abdomen,
and much 3*ellow pollen on the under side of the thorax. There
was also polhn on the chin, and, it may be presumed, on the
proboscis, but this was difficult to observe. I had, however,
1 All from Darwin, Forms of Flowers, 137-1 17, &c.
ADAPTATIONS FOR INTERCROSSING. 239
independent proof that pollen is carried on the proboscis ; for a
small branch of a protected short-styled plant (which produced
spontaneously only two capsules) was accidentally left during
several days pressing against the net, and bees were seen insert-
ing their proboscides through the meshes, and in consequence
numerous capsules were formed on this one small branch. . . .
It must not, however, be supposed that the bees do not get more
or less dusted all over with the several kinds of pollen ; for this
could be seen to occur with the green pollen from the longest
stamens. . . . Hence insects, and chiefly bees, act both as
general carriers of pollen, and as special carriers of the right
sort."
429. Finally, a long series of experiments (requiring eighteen
distinct kinds of union) proved that both kinds of pollen are
nearly or quite impotent upon the stigma of the same flower, and
that no ovary is fully fertilizable by other than a " legitimate
union," i. e. by stamens of the corresponding length ; but that
the mid-length pistil is more prolific than either of the others
under illegitimate union of either kind ; which might perhaps be
expected, as the pollen proper to it is intermediate in size of
grains between that of the long and that of the shortest stamens.
430. Nesaea verticillata, a common Lythraceous plant of the
Atlantic United States, is similarly trimorphous, but has not
yet been particularly investigated. Several South African and
American species of Oxalis are equally trimorphous, and have
been investigated by Darwin and Hildebrand, 1 with results
quite as decisive as in Lythrum Salicaria. One genus of
Monocot}'ledons has trimorphous blossoms, viz. Pontederia, of
which the North American P. cordata is a good illustration. 2
431. All known flowers exhibiting reciprocal dimorphism or
trimorphism are entomophilons : no such wind-fertilized species
is known. Few of them are irregular, and none very irregular :
they do not occur, for instance, in Leguminosse, Labiatae,
1 Monatsber. Akad. Berlin, 1866 ; Bot. Zeit. 1871, &c. According to
Darwin, Fritz Mueller "has seen in Brazil a large field, many acres in extent,
covered with the red blossoms of one form [of an Oxalis] alone, and these
did not produce a single seed. His own land is covered with the short-styled
form of another species, and this is equally sterile ; but, when the three
forms were planted near together in his garden, they seeded freely." Forms
of Flowers, 180.
2 Detected by W. H. Leggett. See Bulletin of Torrey Bot. Club, vi. 62,
170; and for the original discovery in Brazilian species, by Fritz Mueller,
see Darwin's Forms of Flowers, 183, &c. Pontederia has three lengths of
style and counterpart stamens, as in Lythrum Salicaria, each flower having
two sets of stamens, three in each set.
240 THE FLOWER.
Scrophulariacese, Orchidaceae, &c. Nature is not prodigal, and
does not endow with needless adaptations flowers which are
otherwise provided for.
3. ADAPTATIONS FOR CLOSE FERTILIZATION.
432. Even where cross-fertilization in bisexual flowers is
obviously arranged for, it is apt to be tempered with more or
less of close-fertilization. The more exquisite the arrangements
for the former are, the more completely is the plant dependent
upon insect visitation. Failure to intercross is a remote and
small evil compared with failure to set seed at all. In order
therefore that the plan of cross-fertilization may not defeat even
its own end, through too absolute dependence on precarious
assistance, some opportunity for self-fertilization will usually be
advantageous. Also there is a long array of insect-visited
flowers, especially polyandrous ones, in which close fertilization
must be much the commoner result, except where the pollen of
another but wholly similar flower has greater potency.
433. Subsidiary self-fertilization is secured in a great variety
of ways. In Gentiana Andrewsii, which is proterandrous, and
usually cross-fertilized by humble-bees entering bodily into the
corolla, an exposed surface of pollen long remains fresh upon
the ring of anthers girding the base of the style : when the stigmas
separate, they remain for some days simply divergent, but they
at length become so revolute that the receptive surface is brought
into contact with the ring of pollen below. The opening and
closing of blossoms by day or night, the growth of style, fila-
ments, or corolla after anthesis commences, or other changes of
position, may secure a certain amount of self-fertilization in a
subsidiary or even in a regular way. Then certain species, such
as Chick weed, which blossom through a long season, close-
fertilize even in the bud in early spring, when insects are scarce,
but are habitually intercrossed by insects in summer. Somewhat
similarly, according to Hermann Mueller, 1 certain species, such
as Enphrasia officinalis and Rhinanthus Crista-galli, habitually
produce two kinds of blossoms, one larger and more showy,
usually affecting sunny localities, and with parts adapted to
intercrossing by insects ; the other smaller or inconspicuous, and
with anthers adjusted for giving pollen to the adjacent stigma
without aid. There are gradations between these last arrange-
ments, and the more special and remarkable one of dimorphism
with
1 Bcfruchtung der Blumcri durch Insektcn, 294; Nature, viii. 433.
ADAPTATIONS FOR CLOSE FERTILIZATION. 241
434. Cleistogamy. Here the intention and the accomplishment
of self-fertilization are unmistakable. This peculiar dimorphism
consists iii the production of very small or inconspicuous and
closed flowers, necessarily self- fertilized and fully fertile, in
addition to ordiiMny, conspicuous, and much less fertile, though
perfect flowers. Two cases were known to Linnaeus, 1 and one
of them to Dillenius before him ; those of Viola have long been
familiar in the acaulescent species ; Adrien Jussieu made out
the structure of the cleistogamous flowers in certain Malpighiaceae
in 1832, and recorded in 1843 that Adolphe Brongniart had well
investigated those of Specularia, and that Weddel had discov-
ered them in Impatiens Nolitangere. A full account of the then
known cases was given by MohlMn 1863; but D. Mueller, of
Upsala, who examined Viola canina, is said by Darwin to have
given, 3 in 1857, " the first full and satisfactory account of any
cleistogamic flower." The appropriate name of cleistogamous
was given by Kuhn, 4 in 1867, and is now in common use.
435. Cleistogamous flowers are now known in about 60 genera,
of between twenty and thirty natural orders, of very various
relationship, though all but five are Dicotyledons. All but the
Grasses 6 and Juncus are entomophilous as to the ordinary
flowers, and most of these such as have special arrangements for
their intercrossing, either by dichogam}', heterogone dimorphism
or trimorphism (in Oxalis), or such special contrivances as those
of Orchids.
436. It has been said that the ordinary flowers in such plants
are sterile, and perhaps they always are so except when cross-
fertilized : in most cases they are habitually infertile or spar-
ingly fertile. Probably they suffice to secure in every few
generations such benefit as a cross ma}' give, while the principal
1 Campanula (now Specularia) perfoliata and Ruellia clandestina, the
latter a cleistogamous state of R. tuberosa. Linnaeus did not make out the
structure of the flowers, but supposed them to want the stamens.
2 In Bot. Zeitung, xxi. 309.
8 In Bot. Zeitung, xvi. 730.
4 Ibid. xxv. 65. The name ( denoting " closed up " union or fertilization ) has
been written cleistoyc.no'ts, which is not so proper. We prefer deistor/amous to
cleistoyamic (and so of similar terms), as best harmonizing witli the Latin
adjective form, both in form of termination and in euphoniously taking the
accent upon the antepenult.
6 Amphicarpum (Milium amphicarpon, Pursh) is the earliest recognized
cleistogamous Grass, except perhaps Leersia oryzoides. Some species of
Sporobolus are like the latter, and Mr. 0. G. Pringle has recently detected
suc-li flowers concealed at the base of the sheaths in Danthonia. Amer.
Jour. Sci. January, 1878, 71.
16
242 THE FLOWER.
increase is by cleistogamous self-fertilization, which thus offsets
the incidental disadvantage of the former mode.
437. In general, the cleistogamous are like unto the ordinary
flowers arrested in development, some arrested in the almost
fully formed bud, most at an earlier stage, and in the best
marked cases with considerable adaptive modification. In
these, "their petals are rudimentar}' or quite aborted; their
stamens are often reduced in number, with anthers of very small
size, containing few pollen-grains, which have remarkably thin
transparent coats, and generally emit their tubes while still
enclosed within the anther-cells ; and, lastly, the pistil is much
reduced in size, with the stigma in some cases hardly at all
developed. These flowers do not secrete nectar or emit any
odor : from their small size, as well as from the corolla being
rudimentary, they are singularly inconspicuous. Consequently,
insects do not visit them ; nor, if they did, could they find an
entrance. Such flowers are therefore invariably self-fertilized ;
yet they produce an abundance of seed. In several cases, the
3 T oung capsules bury themselves beneath the ground, and the
seeds are there matured. These flowers are developed before,
or after, or simultaneously with the perfect ones." 1 In Grasses,
however, as in some Dicotyledons, there is much less modifica-
tion and more transition. For when Leersia half protrudes its
panicle, in the usual wa}*, the included half is fertile and the
expanded portion sterile (or almost alwaj's so), although the
flowers may open and exhibit well-developed anthers, ovaries,
and stigmas. But when similar panicles remain enclosed in the
leaf-sheaths, they are mostly fruitful throughout.
438. Fully to apprehend the econom^y of cleistogam}' in pollen-
saving alone, and contrariwise to estimate the expense of
intercrossing, one should compare the small number of pollen-
grains which so completely serve the purpose in a typical cleis-
togamous flower (say 400 in Oxalis Acetosella, 250 in Impatiens,
100 in some Violets) with the several thousands of all entomo-
philous cross-fertilized flowers, rising to over three and a half
millions in the flower of a Peony, also their still greater number
in many anemophilous blossoms. To this loss should be added
the cost of a corolla and its action, also of the production of
odorous material and of nectar. No species is altogether cleis-
togamous. Thus cleistogamy, with all its special advantage,
testifies to the value of intercrossing.
1 Darwin, Forms of Flowers, 310.
PERIANTH, OR FLOWER-LEAVES. 243
SECTION V. THE PERIANTH, 1 OR CALYX AND COROLLA IN
PARTICULAR.
439. The distribution of the floral leaves around the axis,
which belongs to phyliotaxy, and their particular disposition in
the bud (estivation), have already been considered in Chap. IV.
Sect. I., II. And most of the morphology of calyx and corolla
has been outlined in the preceding sections of the present chap-
ter. What remains chiefly relates to particulars of form and
to terrain olog3 r .
440. Duration. The differences in this respect give rise to a
few terms, such as the following. Calyx or corolla may be
Persistent, not cast off after anthesis, but remaining unwithered
until the fruit is formed or matured ; as the calj-x in Labiatae, in
Physalis, and most Roses.
Marcetctnt, withering or drying without falling away ; as the
corolla of Heaths, Drosera, &c.
Deciduous, falling after anthesis and before fructification ; as
the petals of Roses, the calyx and corolla of Columbine.
Ephemeral or Fugacious, lasting for only a day ; as the petals
of Poppy, Helianthemum, Purslane, and Spiderwort. In the
two former, the} 7 are cast or earty deciduous, the anthesis lasting
but a day : in the two latter, the anthesis is equally or more
brief, but the petals deliquesce or decay at once without falling,
as does the whole flower of Cereus grandiflorus and other night-
blooming Cactacese.
Caducous, falling when the blossom opens ; as the calyx of
Poppy and Baneberry.
441. Numerical Terms, succinctly denoting the number of
leaves, either of the perianth as a whole, or of any one of its
circles, are common in descriptive botany. The most general
are those which simply specify the number of component leaves,
by prefixing Greek numerals to the Greek name of leaves, ex-
pressing them in Latin form, or transferring them to the Eng-
lish. Thus
Diphyllous, of two leaves (sepals or petals) ; Triphyllous, of
three ; Tctraphyl/ous,offouT ; Pent aphyllous, of five ; Hexophyllous^
of six, and so on. A tulip and a Tradescantia flower have a
hexaphyllous perianth, but composed of two circles, answering
to calyx and corolla ; each TriphyUout.* When the character
* Perlanthium, alias Periyone or Periyonium. (296.)
2 As elsewhere explained, when numerical composition is indicated without
reference to nature of parts, the terms dimerous, trimerous, tetramerous, penta-
merous, &c., may be used. (322.)
244 THE FLOWER.
of the organ, i. e. whether caryx or corolla is to be specified,
the word sepal or petal is employed in the combination ; as,
Disepalou*, of two sepals ; Trisepalous, of three ; Tetrasepalous,
of four ; Pentasepalous, of five (also written 5-sepalous, and ac-
cordingly 2-sepaious, 3-sepalous) , and so on : also,
Dipetalous, Tripetalous, Tt-trapetalous, Pentapetalous (2-5-
petalous), &c., when the corolla is concerned.
442. Monophy 'lions, Monosepalous, and Monopetalous are the
proper terms for perianth (calyx, corolla, &c.) composed of a
single leaf. Likewise Polyphyllous, Polysepalous, and Potypetal-
ous for the case of a considerable but unspecified number of
members. Unfortunately, in the Linnaean and long-prevalent
use, monopetalous was the term employed to designate a corolla
of one piece in the sense, or the fact, of a coalescence or grow-
ing together of two, three, five, or more petals into a cup or
tube ; and so of a calyx, of a whorl of bracts, &c. And poly-
petalous, polysepalous, and polyphjilous were the counterparts
of this, meaning of more than one distinct piece, whatever the
number. The misleading use, consecrated by long prescription,
is not yet abandoned, but will in time be obsolete. In present
descriptive botany, a polyphyllous calyx, or a polypetalous
corolla, or a 5-petalous corolla, would be taken to mean that the
sepals or petals (as the case may be) were distinct or uncom-
bined, and a monopetalous corolla to be one with petals combined
b}' coalescence. (329.)
443. Terms of Union or Separation. The proper term for a
corolla or a calyx the leaves of which are more or less coalescent
into a cup or tube is
Gumopef.alous for such a corolla, Gumosepalnns for the calyx ;
these terms meaning united petals or sepals. The older and mis-
leading names Monopetalous and Monosepalous. although current
up to a recent day, should be discontinued. Another term is
not rarely used in Germany, that of Sympetalous, for the gamo-
petalous (or formerly monopetalous) corolla, therefore Syn-
S'p Ordinarily the sus-
pensor soon disappears.
^ i >s Httacht'd to the ra-
dicular end of the em-
bryo, which consequently alwa}'s points to the foramen or
micropyle of the seed. The process in Gymnosperms is more
complex, and has to be separately described.
533. Polyeinbryony ', the production of two or more emb^os in
one seed, is not uncommon in Gymnosperms (there being a kind
of provision for it), and is of occasional but abnormal occurrence
in Angiosperms, in the seed of Mistletoe, Santalum, &c. In
these it results from the production and fertilization of more
than one embryonal vesicle. Strasburger has recently ascer-
tained that the commoner polyembryony in the seeds of Onions,
Oranges, Funkia, &c., results from the production of adventlve
embryos, which originate in the nucleus outside of the embryo-
sac and wholly independent of fertilization. 1 Two kinds of
1 Strasburser, Ueber Polyembryonie, in Zeitschr. Natunvis. Jena, xii.
1878 (see Amer. Jour. Sci. April, 1879). It was found that when, by exclu-
sion of pollen, the formation of a normal embryo was prevented, no adventive
FIG. 599. Diagram of the suspensor and incipient embryo at its extremity. 600.
The same, with the embryo a little more developed. G01. The same, more developed
still, the cotyledons faintly indicated at the lower end. 602. Same, with the incipient
cotyledons more manifest. 603. The embryo nearly completed.
FIG 604-600. Forming embryo from a half-grown seed of Buckwheat, in three stages.
607. Same, with the cotyledons fully developed.
THE FRUIT. 2G5
anomalous reproduction are therefore now known, which are
intermediate between sexual and non-sexual, between budding
and fruiting propagation, viz.,
Apogamy, which is budding growth or prolification in place of
that which should subserve sexual reproduction. This was die-
covered in Ferns by Prof. Farlow, while a pupil of De Bary, by
whom our knowledge of the process has recently been extended,
and this name imposed. 1 The production of bulblets in place of
seed or embryo answers to this in Flowering plants.
Parthenogeny , the counterpart analogue of apogamy, is the
non-sexual origination of an embryo extraneous to the embryonal
vesicle or even the embiyo-sac. However abnormal, its occur-
rence is probably not so rare as has been supposed.
CHAPTER VII.
THE FRUIT.
SECTION I. ITS STRUCTURE, TRANSFORMATIONS, AND DEHISCENCE.
534. The Fruit consists of the matured pistil or g3'no2cium
(as the case may be) , including also whatsoever may be joined
to it. It is a somewhat loose and multifarious term, applicable
alike to a matured ovary, to a cluster of such ovaries, at least
when somewhat coherent, to a ripened ovary with calyx and
other floral parts adnate to it, and even to a ripened inflores-
cence when the parts are consolidated or compacted. Fruits,
accordingly, are of various degrees of simplicity or complexity,
and should be first studied in the simpler forms, namely, those
which have resulted from a single pistil. Such a fruit consists
of Pericarp with whatever may be contained in it and incorpo-
rated with it.
embryo appeared in those seeds which habitually produce them. To this
Caelebogync offers an exception. The female of this dioecious plant habit-
ually matures fertile seeds, with a well-formed embryo, in Europe when there
are no male plants in the country. Strasburger ascertained that the embryo
thus formed is adventive, the embryonal vesicle perishing. Parthenogenesis,
of which Caelebogyne was the most unequivocal case, is thus confirmed, and
is shown to occur in most polyembryony ; but it is at the same time explained
to be a kind of prolification.
1 See Farlow, in Proc. Am. Acad. ix. 68; De Bary, Bot. Zeit. xxxvi. 465-487.
286
THE FKU1T.
535. The Pericarp, or Seed-vessel, is the ripened ovary. It
should, therefore, accord in structure with the ovary from which
it is derived. Yet alterations sometimes take place during fruc-
tification, either by the abortion or obliteration of parts, or by
accessory growth.
536. Internal Alterations. Thus, the ovary of the Oak con-
sists of three cells, with a pair of ovules in each ; but the fruit
has a single cell, filled with a solitary seed, onty one ovule being
matured, while two cells and five ovules are suppressed, the
remains of which may be detected in the acorn. The ovary
of the Chestnut has six or seven cells, and a pair of suspended
ovules in each ; but only one of the dozen or fourteen ovules
ever develops into a seed, except as a rare monstrosity. The
three-celled ova^ of the Horsechestnut and Buckeye is similar
in structure (Fig. 608-611), and seldom
ripens more than one or two seeds ; but the
abortive seeds and cells are obvious in the ripe fruit. The
ovaiy of the Birch and of the Elm is two-celled, with a single
ovule in each cell : the fruit is one-celled, with a solitary seed ;
one of the ovules being uniformly abortive, while the other in
enlarging thrusts the dissepiment to one side, and obliterates the
empty cell. Similar suppressions in the fruit of parts actually
extant in the ovary are not uncommon.
537. On the other hand, there may be more cells in the fruit
than there are primarily in the ovary. Thus the fruit of Datura
J3 dicarpellary and normally two-celled, with a large placenta
projecting from the axis far into the cells. But each cell be-
comes bilocellate, that is, divided into two, by a false partition
growing out from the back of each carpel and cohering with the
middle of the adjacent placenta. So the 5-carpellary and nor-
mally five-celled ovary of common Flax early becomes spuriously
ten-celled (morphologically speaking, not 10-locular, but 10-
locellate), by a false partition extending from the back of each
FIG. 608. Longitudinal section of the ovary of a Buckeye (^Esculus Pavia), showing
the pairs of ovules in two of the cells. fi()9. Transverse section of the same displaying all
three cells and six ovules. 610. Same of half-grown fruit, with single fertile seed, abor-
tive ovules and obliterating cells. 611. Dehiscent one-seeded fruit, diminished in size.
ITS STRUCTURE AND TRANSFORMATIONS. 287
carpel across its cell (Fig. 539-541) ; and the solitary carpel
is similarly divided lengthwise in many species of Astragalus,
as in Fig. 534. Transverse divisions or constrictions across
a maturing ovar} T (such as is seen in Fig. 620) are not uncom-
mon, especially in legumes and other pods, and are of little mor-
phological significance.
538. External Accessions may here be referred to. The wing
of the pericarp in Maple, Ash, and the like (Fig. 625-627), are
familiar instances of this ; and of the same nature are the im-
bricated scales which cover some Palm-fruits ; the prickles on
the pod of Datura, Ricinus, &c., and the hooked or barbed
prickles of many small pericarps (as in various Borraginacese),
which thus become burs and are disseminated by adhering to the
haiiy coat of cattle. All these are of the nature of superficial
outgrowths, and these especially affect the pericarp or parts
connected with it.
539. Persistence of Connected Organs. An adnate calyx (331) ,
being consolidated with the ovary, necessarily makes a constit-
uent part of the fruit, in the pome (575) doubtless a very large
part. The limb or lobes of such adnate organ may persist, as
the tips of the sepals on an apple or quince, and may be turned
to useful account, as is the pappus of Composite for dissemina-
tion. Or, in small pericarps, the style may persist as part of the
fruit, and subserve the same ends, either by becoming feathery
for aerial dissemination, as in Clematis and in one section of
Geum, or by becoming hooked at the tip for adhesion to fleece,
&c., as in other species of the latter genus. Or adjacent parts
which are not actually incorporated with the pericarp may play
similar parts in the economy, as the hooks on the calyx-tube of
the dry calyx of Agrimonia, which at maturity is detached with
the included fruit, the fleshy fructiferous calyx of Gaultheria (Fig.
651) and of Mulberry (Fig. 654) ; and the pulp}' fructiferous re-
ceptacle of the strawberry (Fig. 653) : the ultimate utilities in
both classes of instances being similar, viz., wide dispersion of
the seed by animals, whether by external carriage, or b} T being
devoured and the voided seeds of fleshy fruits thus disseminated.
540. Transformations in Consistence. In the change from
ovary to mature pericarp, various kinds of transformations may
take place. In some the wall of the ovary remains thin and
becomes in fruit foliaceous or leaf-like, as in a pea-pod, the
carpels of Columbine, and Marsh Marigold (Caltha) , or the pod
of Colutea or Bladder Senna. In others it thickens and becomes
at maturity either dry throughout, as in nuts and capsules ; or
fleshy or pulpy throughout, as in berries ; or hard-rinded with-
288 THE FKUIT.
out but soft within, as in a pepo ; or flesh}' or beny-like without,
but indurated within, as in all stone-fruits, such as the cherry
and peach.
541. When the walls of a pericarp consist of two layers of dis-
similar texture (as in a peach) the outer layer is called EXOCARP,
the inner ENDOCARP, these terms meaning exterior and interior
parts of a fruit. When the external layer is a comparatively thin
stratum or film, it is sometimes termed the EPICARP. When it is
fleshy or pulpy it is named SARCOCARP. When the endocarp within
a sarcocarp is hard and bony or crustaceous, forming a shell or
stone, this is termed a PUT AMEN. When three concentric layers are
distinguishable in a pericarp, the middle one is called MESOCARP.
542. Fruits may be divided into two kinds, in reference to
their discharging or retaining the contained seeds. They are
dehiscent when they open regularly to this end ; indehiscent when
they remain closed. There is a somewhat intermediate condi-
tion, when the}' rupture or burst irregularly, as in Datura Metel,
&c. Dry pericarps with single seeds are commonl}* indehiscent ;
those with several or many seeds mostly dehiscent. Seeds pro-
vided with a wing or coma or any analogous help to dispersion
are always in indehiscent pericarps. Permanently fleshy peri-
carps are indehiscent, stone-fruits as well as berries. But in
some stone-fruits (/. e., with indurated endocarp and fleshy
exocarp), such as those of Almond (Fig. 640) and Hickory,
the barely fleshy exocarp or sarcocarp dries or hardens, instead
of softening, as maturity is approached, and at length separates
from the putamen b}* dehiscence.
543. Dehiscence, the opening of a pericarp for the discharge
of the contained seeds, is regular or irregular ; or, better, is
normal and abnormal. For most of the abnormal or non-t3*pical
modes are as determinate and uniform in occurrence as the typi-
cal modes. A good English name for dehiscent pericarps in
general is that of POD.
544. Regular or normal dehiscence is that in which a pericarp
splits vertically, for its whole or a pail of its length, on lines
which answer to sutures or junctions, that is, along lines which
correspond to the margins or midribs of carpellary leaves, or to
the lines and surfaces (or commissures) of coalescence of con-
tiguous carpels. The pieces into which a pericarp is thus sun-
dered are termed VALVES.
545. The normal dehiscence of a carpel is by its inner, ven-
tral, or ovuliferous suture, that is, by the disjunction of the
leaf-margins, as in Fig. 618. Its only other line of normal
dehiscence is by the opposite or dorsal suture, that is, down
DEHISCENCE.
289
the midrib. Legumes usually dehisce by both sutures (as in
Fig. 619), therefore into two valves.
546. A dehiscent pericarp formed of two or more carpels is
called a CAPSULE. The two leading terms descriptive of capsular
dehiscence were based upon the modes of opening of pericarps
having as many cells as carpels : they are the septicidal, that is,
as the term denotes, cutting through the septa or dissepiments ;
and the loculicidal, that is, cutting into the Loculi or cells.
547. Septicidal, the dehiscence through the dissepiments, is
the disjunction of a pericarp into its constituent carpels, these
then usually themselves dehiscing down their ventral suture,
as in Fig. 612, illustrated by
the diagram, Fig. 613. Good
examples are furnished by the
Hypericum Family (the pistil
illustrated in Fig. 536, 537),
where the placentae which
compose the axis are carried
awa} r on the edges of the par-
titions or introflexed valves ;
also by Rhododendron, Kal-
mia, and the like, in which
the placentae remain combined
into a column in the axis (the
COLUMELLA or column) , from
which the edges of the valves
break awa} r .
/ \ 548. The septicidal dis-
* junction of the carpels does
not of itself open the cells.
Such separated carpels when
one-seeded not rarely remain
closed, as in Mallow, Ver-
bena, &c. Or when dehiscent they may open both by the ventral
and dorsal sutures ; i. e. , the pericarp may first divide into its
constituent carpels, and then each carpel break up into half
carpels, as in Euphorbia.
549. Loculicidal, the dehiscence into the loculaments, loculi,
or cells of the pericarp (shown in Fig. 614, and the diagram,
615), is that in which each component carpel splits down its
/N
FIG. 612. Septicidally dehiscent tricarpellary capsule of Elodes Virginica. 613. Dia-
gram of septicidal dehiscence.
FIG. 614. Loculicidally dehiscent tricarpellary capsule of an Iris, divided trans-
versely at the middle. 015. Diagram of loculicidal dehiscence.
290 THE FRUIT.
dorsal suture, as in Iris, Hibiscus, CEnothera, &c. In this, the
dissepiments remain intact. If the}' break away from the centre
then the}* are borne on the middle of the valves, as in the figures
above cited. If they remain coherent in the axis but break away
from the valves, the result is one form of what is called
550. Septifragal dehiscence, i. e., a breaking away of the
valves from the septa or partitions, as shown in Fig. 616. This
represents the loculicidal form of the septifragal mode, which is
less common than that of the accompanying diagram, Fig. 617.
Here the partitions alternate
s^****^^ with the valves ; that is, the
dehiscence of the pericarp is
/ I l ^^^^ \ of the septicidal order, as
I ^^\v^ I I ) near as may be, but the par-
V J V J titions do not split, wherefore
^>^ *S \^ ^/ the valves break away at the
1516 6I7 common junction. To this
the term marginicidal has been applied. It occurs in the 2-3-
carpellary capsule of Ipomaea (especially in the common Morning
Gloiy) , in the 5-carpellary capsule of the North American species
of Bergia ; likewise in the 2-carpellary pod of Cruciferae (Fig.
623), with a difference that the placentae from which the valves
break away are here parietal and the partition is abnormal.
551. The terms septicidal and loculicidal apply equally in plan,
though not with etymological correctness, to one-celled capsules
with either parietal (495) or free central (599) placentae. When
the dehiscence is of the septicidal type and the placentation pari-
etal, the (half) placentae are borne on the margin of the valves,
as in the Gentian family and the species of Hypericum with one-
celled capsule. When the placentae are borne on the middle
of the valves, as in Violets, the dehiscence is of the loculicidal
type. In the case of free central placentae with no trace of
partitions, the character of the dehiscence may usually be deter-
mined by the position of the styles or stigmas relative to the
valves.
552. Dehiscence may be quite normal although very partial,
as when confined to the apex of the capsule of Cerastium and
of Primula, and even to the pores under the radiate stigmas
of Poppy.
553. Irregular or abnormal dehiscence is such as has no respect
to the normal sutures ; as where the dehiscence is transverse ;
FIG. 616. Diagram of loculicidally septifragal dehiscence. 617. Same of septicidally
or rather marginicidally septifragal dehiscence.
ITS KINDS. 291
either extending part waj r round, as in the pod of JcfTcrsonia,
or completely round, so that the upper part falls off like an
unhinged lid. This circumscissile dehiscence occurs in many
plants of widely different orders ; such, for example, as Purslane
(Fig. 621), genuine Amaranths, Plantain, Pimpernel, and Hen-
bane. In other cases, as in Antirrhinum (Snap-dragon) and
its allies, the cells burst by irregular laceration at a definite
point, and discharge the seeds through the ragged perforation ;
or one or more neat valvular orifices are formed on some parts
of the wall, as in Campanula.
SECTION II. THE KINDS OF FRUIT.
554. FRUITS have been minutely classified and named; 1 but
the terms in ordinary use are not very numerous. A rigorously
exact and particular classification, discriminating between the
fruits derived from simple and from compound pistils, or between
those with and without an adnate calyx, is too recondite and
technical, and sometimes too hypothetical, for practical pur-
poses. It is neither convenient nor philosophical to give a
substantive name to every modification of the same organ. For
all ordinary purposes, both of morphological and systematic
botany, it will suffice to characterize the principal kinds under
the four classes of
Simple fruits, those which result from the ripening of a single
pistil ;
Aggregate, those of a cluster of carpels of one flower crowded
into a mass ;
Accessory or Anthocarpus, where the principal mass consists
of the surroundings or support of either a simple or an aggregate
fruit ;
Multiple or Collective, formed by the union or compact aggre-
gation of the pistils of several flowers, or of more than one.
555. Simple Fruits may be distinguished, upon differences of
texture, into Dry Fruits, Stone Fruits, and Baccate Fruits ; or,
better, into Dry and Fleshy ; and the first may be divided into
1 The greater part of the forty-three substantive names of Desvaux's,
and even of the thirty-six of Dumortier's and of Lindley's elaborate classi-
fications of fruits have never found employment in systematic botany, and
doubtless never will be used. Yet a detailed carpological classification has
its uses for the student. Among the more recent attempts are the successive
ones of Dickson, McNab, and Masters. See Nature, iv. 347 (also in Trimen's
Jour. Bot. 1871, 310), iv. 475, and v. 6.
292
THE
dehiscent and indehiscent kinds. 1 Theoretically, each kind may
be divided into those of a simple and those of a compound pistil,
and some would make the primary division on this character.
Some also would separate fruits with adnate or superior calyx
from those free of all such combination. But in practice these
differences can seldom be indicated by substantative names.
The name of berry is equally applicable to the fruit resulting
from the single carpel of Actaea, the syncarpous ovary of the
grape, and the similar ovaiy with adnate calyx of a gooseberry
and cranbeny. It should be understood that the kinds shade
off one into another most freely.
556. Dehiscent Fruits (543), or Pods, are distinguishable into
apocarpous, or of single carpels, and syncarpous, of more than
one carpel, i. e. the first of a simple, the second of a compound
pistil. The first kind is mainly represented by the Follicle and
the Legume ; the second, b} T the Capsule and its modifications.
557. A Follicle is a pod formed
of a simple pistil, and dehiscent by
one suture (this almost always
the ventral or inner suture) alone ;
as in the Larkspur, Columbine,
Peony, and Marsh-Marigold (Fig.
018 618) ; also in Milkweed and Dog-
bane. There ma}* be several follicles or only
one to a flower, even in the same genus, as in
Larkspurs, Cimicifnga, &c. In Magnolia
(Fig. 648-650), fleshy carpels become follicles
dehiscent by the dorsal suture.
558. A Legume is the pod formed of a
simple pistil which is dehiscent by both sut-
ures (as in the Pea, Fig. 619), so dividing
into two pieces or valves. (544.) This is the
fruit of the Pulse Family, accordingly named
Lcguminosae (Leguminous plants) : indeed,
the name of legume is restricted to the fruits of this family,
and in descriptive botany is extended to all the modifications
1 Dr. Masters's modification of Dickson's and McNab's classification of
simple fruits, as to primary kinds, is into
1. Nuts, or Achccrocarps, dry and indehiscent ;
2. Pods, or Rcgmacarps, dry, dehiscent ;
3. Stone-fruits, orPyrcnocarps, fleshy without, indurated within, indehiscent ;
4. Berries, or Sarcocarps, fleshy throughout, indehiscent.
FIG. 618. A dehiscent follicle of Marsh-Marigold, Caltha palustris.
FIG. 619. Legumo cf a Sweet Pea, already dehiscent. 620. Loment of a Desmodium.
ITS KINDS.
293
which that order presents. Some of these, in fact, are in-
dehiscent and reduced to akenes ; some break up at maturity
into one-seeded indehiscent articulations or joints, which are
dispersed as if they were so many seeds. A legume of the latter
kind takes the special name of LOMENT, Lat. Lomentum. (Fig.
620.) In Mimosa (Sensitive-plant, &c.), such articulations de-
hisce into two valves. They also fall away from the sutures,
or from a persistent marginal border of them, or in some cases
the valves thus fall away entire. The persistent frame which
remains has been called a REPLUM, an architectural word, here
taken in the sense of door-case.
559. A Capsule is the pod, or dehiscent fruit, of any compound
pistil. When regularly and com-
pletely dehiscent, as already stated
(544), the pod splits lengthwise into
pieces or valces. The modes of regular
dehiscence are illus-
trated in Fig. 612-
617. Two modifica-
tions of the capsule
have received distinc-
tive names which are
in common use, viz.
the Pyxis and the
Silfquc.
560. A Pyxis or Pyxidium is a dry fruit which opens by a
circular line, cutting off the upper part as a lid ; *. e., the dehiscence
is circumscissile. (553, Fig. 621.) In the Purslane,
Pimpernel, Henbane, and Plantain, the pyxis is
a capsule ; in Amaranths (Fig. 637) it is a
utricle ; in Jeffersonia (Fig. 622) it is a modi-
fication of the follicle, being of one carpel which
dehisces transversely, and not all round, so that
the lid remains attached. 6 - 4
561. A Silique is a narrow two-valved capsule, with two pari-
etal placentae, from which the valves separate in dehiscence ; as
in plants of the Cruciferous or Mustard family (Fig. 623),
to the fruit of which this term is restricted. Usually, a false
partition is stretched across between the two placentae, render-
FIG. 621. Pyxis of Purslane, Portulaca oleracea, the top separating entirely and
falling away.
FIG. 622. Pyxis-like follicular fruit of Jeffersonia diphylla; the lid remaining
attached dorsally.
FIG. 623. SiiiqueofCardamine in dehiscence. 624. Silicleof Capsellaor Shepherd's-
Purse, lateral view, and an oblique view of the same with one valve removed.
294
THE FKUIT.
ing the pod two-celled in an anomalous manner. A SILICLE
(Sitictda, diminutive of siliqua) is merely a short silique, the
length of which does not more than twice or thrice surpass the
breadth; such as that of Shepherd' s-Purse (Fig. 624). and of
Lunaria, Candytuft, &c.
562. Indehiscent Dry Fruits are almost always one-seeded or
very lew-seeded. If numerous, the seeds thus placed would not
be dispersed. The ordinary kinds are strictly one-
seeded, and in common language are often con-
founded with seeds. The ways in which such fruits
are dispersed are various. In the following case,
the adaptation of the pericarp to dispersion by wind
distinguishes the species of fruit.
563. The Samara, sometimes called in English a
Key, is an indehiscent one-seeded fruit provided
with a wing. In the White Ash
the wing is terminal (Fig. 625) ;
in other species the whole fruit
is wing-margined ; in Birch and
Elm (Fig. 626) the wing sur-
rounds the body of the pericarp ;
and the Maple fruit is a double
samara or pair of such fruits, con-
spicuously winged from the apex.
564. Akene (Lat. Achenium) is a general name for all the
one-seeded, dry and hard, indehiscent and seed-like small fruits,
such as are popularly taken for naked
seeds. But that they are true pistils,
or ovaries ripened, is evident from the
style or stigma they bear, or from the
scar left by its fall ; and a section
brings to view the seed within, provi-
ded with its own proper integuments.
The name has been restricted to the seed-like fruits of simple
pistils, such as those of the Buttercup (Fig. 628, 629) , Anemone,
Clematis, and Geum. The style in some species of the latter
remains on the fruit as a long and feathery tail, in others as a
short and hooked one, both being agents of dissemination. The
grains of the strawberry (Fig. 653) are also akenes. The name
is extended to all one-celled seed-like fruits resulting from a
FIG. 625. Samara or key of White Ash. Fraxinus Americana. 626. That of White
Elm. Ulmus Americana. 627. DonMe samara of lied Maple, Acer rubrum.
FIG. 628. Aclieniuru of a common Buttercup. 629. Vertical section, showing the
seed within.
ITS KINDS.
295
compound ovary, and even when invested with an adnate calyx-
tube. Of the latter is the fruit of Composite. (Fig. 630-635.)
Here the tube of the calyx is incorpo-
rated with the surface of the ovary ;
and its limb or border, obsolete in some
cases (Fig. 630), in others appears
as a crown or cup (Fig. 631), or set of
teeth or of scales (Fig. 632, 633), or as
a tuft of bristles or hairs (Fig. 634,
635), &c., called the PAPPUS. In the Lettuce and Dandelion
(Fig. 635), the achenium is rostrate, or beaked, i. e. its summit
is extended into a slender beak. An akene with adnate ealyx
has been termed a CYPSELA.
565. The Utricle is the same as the akene, only with a thin
and bladder}' loose pericarp, like that of Goosefoot.
(Fig. 636.) This thin coat sometimes bursts irregu-
larly, discharging the seed. In the true Amaranths,
the utricle opens by a circular line, and the upper
part falls as a lid, converting the fruit into a small
pyxis (560), a transition form. (Fig. 637.)
566. A Caryopsis or Grain differs from the utricle
or akene in having the seed completely filling the
cell, and its thin coat firmly consolidated throughout
with the very thin pericarp ; as in wheat, Indian
corn, and all other cereal grains. Of all fruits this
is the kind most likely to be mistaken for a seed.
567. A Nut is a hard, one-celled and one-seeded, indehiscent
fruit, like an achenium, but larger, and usually produced from
an ovary of two or more cells with one or more ovules in each,
all but a single ovule and cell having disappeared during its
growth (536) ; as in the H^zel, Beech, Oak (Fig. 638), Chest-
FIG 630. Arhenium of Mayweed (no pappus). 631. That of Cichory (its pappus a
shallow cup). (532. Of Sunflower (pappus of two deciduous scales). 633. OfSneezeweed
(Hclenium), witli its pappus of five scales. 634. Of Sow-Thistle, with its pappus of
delicate downy hairs. 635. Of the Dandelion, tapering below the pappus into a
long beak.
FIG. 636. Utricle of fhenopodium album, or common Goosefoot. 637. Utricle of an
Amaranth, by transverse dehiscence becoming a pyxis.
296 THE FRUIT.
nut, and the like. The nut is often enclosed or surrounded by
a kind of involucre, termed a Cup tile ; such as the cup at
the base of the acorn, the bur of the chestnut, and the leaf-
like covering of the hazel-nut. The name Glans
(sometimes Gland in English) is technically applied
to such nuts, this being their classical Latin name.
568. The fruit of the Walnuts and Hickory is
apparently a kind of drupaceous nut, or something
intermediate between a stone fruit and a nut. But
certain monstrosities give reason for supposing that
the seeming exocarp (541), which in Hickory
hardens and at maturity dehisces in four valves,
is of the nature of an adnate involucre. The cocoanut is a sort
of fibro-drupaceous nut.
569. Nutlet, or in Latin form NUCULE (Nucula), is sometimes
superfluously employed in a literal sense, as a diminutive nut. 1
Of late it has acquired a good and fairly legitimate use as the
name of the seed-like, or rather akene-like, closed parts or lobes,
of crustaceous or other hard texture, into which certain bilocular
or plurilocular pericarps separate at maturity, i. e. for the seg-
ments of a schizocarp, 571, which resemble akenes. 2 These are
sometimes carpels, sometimes half-carpels, as in Verbena, also
in Borraginacese and Labiatse (in which the segments are greatly
separated in the ovary), and sometimes, as in Nolana, they are
portions of compounded carpels which have been exceedingly
multiplied by chorisis.
570. There are complete transitions between dry nutlets, with
a thin and herbaceous epicarp, and the pyrence (574) or stony
inner portion of such carpels when drupaceous or composing a
drupe of two or more stones. It is therefore a hardly incongru-
ous and ver}' convenient use which extends the term nutlet to
include these small seed-like stones also, as, for example, to
those of Holly, Bearberry, Hawthorn, and the like.
571. The pair of achenium-like or often samara-like carpels,
1 Nut and akene, between which there is no fixed distinction, will cover
this ground. The fruit of Cyperaceae, for instance, is truly an achenium,
if this name is ever to be used (and it now commonly is) for any other than
a monocarpellary fruit. It is often termed a nut, sometimes a nutlet, and
by a late writer, Boeckler, a caryopsis.
2 Cocci (sing. Coccus, from a Greek word for kernel) is another name for
fruit-carpels, or separating lobes of a dry pericarp, as well for dehiscent ones
(of Euphorbia) as for indehiscent. Hence such lobed or partible fruits
are said to be dicoccous, tricoccous, &c., according to the number of lobes or
carpels.
FIG. 638. Acorn (nut) of White Oak, with its cup, or en pule.
ITS KINDS. 297
united by their inner face but separating entire at maturity,
which constitute the fruit of Umbelli ferae, takes the name of
CREMOCARP (Lat. Cremocarpium) ; and the halves are called
MERICARPS. These names it may sometimes be convenient to
use ; yet it is not advisable to have special names for the fruits
of particular families ; and mericarp is here synonymous with
carpel. For dry fruits in general (or such as become dry)
which are composed of two or more carpels, and which at matu-
rity split up or otherwise separate into two or more closed one-
seeded portions, an appropriate recent name is that of SCIIIZOCARP.
The component carpels of such a fruit were long ago named Car-
cerules (carceruli, little prisons) by Mirbel.
572. Fleshy Fruits, which from their texture are naturally
indehiscent, may be either fleshy throughout, or with a firm rind
or shell, or fleshy externally and hard or stony internally. Of
the latter, the type is
573. The Drupe or Stone Fruit proper (Fig. 639), that of the
cherry, plum, and peach. True drupes are of a single carpel,
one-celled and one-seeded
(or at most two-seeded) , in
the ripening of which the
outer portion of the pericarp
becomes fleshy or pulpy, and
the inner stony or crustace-
ous, i. e. divides into sarco-
carp and putamen. (541.)
But the name is extended to
pericarps of similar texture resulting from a compound pistil,
either of a single cell, as in Celtis, and (by abortion) in the olive,
or of two or several cells, as in Cornus, Rhamnus, &c. The several
pericarps of the aggregate blackberry and raspberry are diminu-
tive drupes or DRUPELETS.
574. Small drupes are often confounded with berries, and the
stone or stones taken for seeds. Especially is it so in drupes
or drupaceous fruits of more than one cell, ripening into separate
or separable hard endocarps or stones, each filled b} T a seed. 1
Bearberries (Arctostaplrylos) and Huckleberries (Gaylussacia)
are good illustrations of this. The seed-like endocarps of this
1 The term Acinus, the original name of such a berry as a grape, has heen
used in descriptive botany for a small drupe or drupelet, and the ripened
carpels of Rubus have been termed acini or acines, but without discriminating
them from berries.
FIG. 639. Vertical section of a peach. 640. An almond ; in which the exocarp, the
portion of the pericarp that represents the pulp of the peach, remains juiceless, and at
length separates by dehiscence from the endocarp, cr shell
298
THE FRUIT.
sort are PYREN.E ; and the fruits are dipyrenous, tripyrenous,
tetrapyrenous, &c., according as they contain two, three, or four
p}Tenae. When the sarcocarp is thin and dries up at maturity,
these pyrenae pass by gradations into nuculae (569) or nutlets:
hence pyrenae are not uncommonly in English descriptions called
nutlets or nucules.
575. The Pome (Fig. 641, 642) is the name of the apple, pear,
and quince. These are fleshy fruits, composed of two to
several carpels (rarely by abortion only one) , of
parchment-like or (in Hawthorns) bony texture,
enclosed in flesh which morphologically belongs
to adnate catyx and receptacle ; as may be ap-
prehended b}- comparing a rose-hip (Fig. 407, in
flower) with an apple or a pear. Of the quince, the
whole flesh is calyx or hypanthium (395) ; in the
apple and pear, the inner or core-portion of the
flesh is of the nature of disk, investing the carpels.
In the fruit of Hawthorns, the carpels become bony
pyrenae (574), and so the fruit is drupaceous, is
indeed nothing more than a s}'ncarpous drupe.
In Eriobotrya, or Cumquat, the carpels becoming
very thin and membranaceous, the pomaceous
fruit is in fact a kind of berry.
576. The Pepo, or Gourd-fruit (Fig. 643), of which the gourd
and squash are the type, and the melon and cucumber equally
familiar illustrations, is the char-
acteristic fruit of Cncurbitaceae,
fleshy internally and with a hard
or firm rind, all or part of which
is referable to the adnate calyx
completely incorporate with the
ovary. This is either one-celled
with three broad and revolute
parietal placentae, or these pla-
centae, borne on thin dissepiments,
meet in the axis, enlarge, and
spread, unite with their fellows
on each side, and are reflected to
the walls of the pericarp, next which they bear their ovules. As
the fruit enlarges, the seed-bearing placentae usually cohere
with the walls, and the partitions are obliterated, giving the
FIG. 641. Pome or apple in transverse section. 642. Quince in vertical section: the
inner flesh answering to disk in the apple and pear is here wanting.
FIG. 643. Section of the ovary of the Gourd. 644. Diagram of one of its constituent
carpels.
ITS KINDS.
299
appearance of a peculiar abnormal placentation, which the study
of the ovary readily explains. In the watermelon the edible
pulp all belongs to the greatly developed placentae. Fruits of
this family in which the rind also is soft at maturity are true
berries.
577. The Hesperidium (orange, lemon, and lime) is the fleshy
fruit of a free mairy-celled ovary with a leathery rind, and is a
mere variety of the berry. The name is,
applied only to fruits of the Orange tribe.
578. The Berry (Lat. Bacca) comprises
all simple fruits in which the pericarp is
fleshy throughout. The grape, gooseberry,
currant, cranberry (Fig. 045), banana, and
tomato are familiar examples. The first
and last consist of an ovary free from the
calyx ; in the others, calyx and ovary are
combined by adnation.
579. Aggregate Fruits are those in which
a cluster of carpels, all belonging to one
flower, are crowded on the receptacle into
one mass, as in the raspberry and black-
berry taken as a whole. (Fig. 64G.) They
may be aggregates of an}' kind of simple
fruits. But when diy and not coherent, the mass would simply
and properly be described as a head or spike of carpels, more
commonly of akenes, as in Ranunculus, Ane-
mone, &c. Yet when numerous carpels thus
compacted become flesh}', and sometimes more
or less coherent, the aggregate may need to be
taken into account. The best name for it is
that of SYNCAKPIUM, or in English
form SYXCAKP. But the term lias
been applied to multiple fruits as
well. 1 In Hydrastis, the numerous
carpels imbricated on the upper
part of the torus arc baccate, that
is, become berries ; in araspberiy,
the seemingly baccate grains are drupaceous (being drupelets, 573),
1 The si/ncarp which is a gynoecium might be designated a simple sijn-
rnrpiwn ; that which is an inflorescence, a complex syncarpium, which may be
biflorous, pauciflorous, or multiflorous.
FIG. 645. The larger Cranberry, Vaccinium (Oxycoccus) macrocarpon ; the berry
transversely divided.
FIG. 64r>. Vortical Faction of lialf of r_ blackberry (of Rul-us villosus), enlarged ; aiid,
647, of one <:i its drupelets more ma^nilie;!.
C TT
300
THE FRUIT.
and, slightly cohering together (though without organic union) ,
they fall as one body from the conical dr}' torus at maturity. It
is the same in blackberries or bramble-berries (Fig. 64G, 647),
except that the drupelets persist on the torus, which partakes of
the juiciness. 1 In the aggregate fruit of Magnolia (Fig. 648-650),
such carpels, imbricated over one another, cohere more or less
at all contiguous parts, and
become drupaceous ; never-
theless, at maturity each
opens dorsally, allowing the
seeds to fall out : in age it
dries and hardens, and also
separates from its comu c-
tions, and so be-
comes a follicle, but
with the remark-
able peculiarity of
dorsal instead of
ventral dehiscence.
(Fig. 650.) In Li-
riodendion, a tree
of the same family,
such carpels are
and indehiscent throughout ; and they largely consist of long
and flat st}ies, imbricated in a cone, but separating from each
other and from the slender torus at maturit}', when each becomes
a samara.
580. Accessory or Anthocarpous Fruits are those of which some
conspicuous portion of the fructification neither belongs to the
pistil nor is organically united with it, except by a common
insertion. The part thus imitating a fruit, while it is really no
part of the pericarp, is sometimes called a Pseudocarp, or an
Anthocarp or Anthocarpium. This condition may occur either
in simple, in aggregate, or in multiple fruits.
1 The aggregate fruit like that of Rubus (named by some Conocarpiinn,
by others an JEterio, En/throstomiun, &c.) was termed by Dumortier a Drupe-
turn. A similar aggregation of baccate carpels he termed a Baccetum ; of
follicles, a Follicetum, &c. All such names may look well in a system ; but
they are both superfluous and unmanageable in phytography.
FIG. 648. Aggregate fruit of Umbrella-tree, Magnolia Umbrella, reduced in size; a
seed, from a lower dehiscent carpel hangs on a thread, consisting of a tuft of extensile
spiral ducts unravelled. 649. Same in longitudinal section. 650. One of the carpels
detached, at full maturity, drfed up, dqrsally dehiscent, exposing the pajr. of seedg of
the natural size.
ITS KINDS.
301
581. Gaultheria procumbens, the aromatic Wintergreen (Fig.
651, 652), affords a good example of the first. Its seeming
berry (the checkerberry) , with summit crowned by the tips of
the calyx-lobes, well imitates the true
berry of a Vaccinium, such as that of
Fig. 645. But it comes from a flower
with thin calyx, underneath and free
from the ovary. Its fruit is really
a capsule : in the process of fructi-
fication, the calyx enlarges, becomes
succulent, completely encloses the capsule or true fruit, yet
without adhering to it, and in ripening counterfeits a red
berry. So in Shepherdia, or Buffalo Berry, the seeming sarco-
carp of a drupe is really a free calyx, accrescent and succulent,
enclosing an akene. So, also, the apparent achenium or nut of
Mirabilis, or Four-o'clock, and of its allies,
is the thickened and indurated base of the
tube of a free catyx, which contracts at the
apex and encloses the true pericarp (a utricle
or thin akene) , but does not cohere with it.
582. Likewise the torus, although not con-
spicuous, may be said to be an accessory part
of the aggregate fruit of the Blackbeny or
Bramble (579) : it becomes the solely con-
spicuous and the sole edible part of a straw-
berry (389, Fig. 406, 653), the akenes or
true fruits dispersed over the surface being
apparently insignificant. Equally in man} T
multiple fruits the conspicuous flesh belongs to receptacle (either
torus or rhachis), to calyx, or even in part to bracts, or to all
these parts combined, as in a pine-apple.
583. Multiple or Collective Fruits 1 are those which result from
the aggregation of several flowers into one mass. The simplest
of these are those of the Partridge-Berry (Mitchella, Fig. 467),
r
1 Collective is the preferable name. The term multiple was applied by
DeCandolle to what are here (following Lindley) called aggregate, fruits ;
and the aggregate fruits of DeCandolle are here called multiple or collective.
Moreover, the distinction between accessory or anthocarpous and collective
or multiple fruits was not recognized by Lindley, who combined the two
in his original "Introduction to Botany." In this work four classes are
given : 1. Fruit simple, APOCARPI ; 2. Fruit aggregate, AGGREGATI ;
FIG. 651. Forming capsule of Gaultheria procumbens, with enlarging calyx partly
covering it. 652. Same, more advanced, and in longitudinal section.
FIG. 653. Vertical section of half a strawberry. Compare with Fig. 406.
302
THE FKUIT.
and of certain species of Hone} r suckle, formed of the ovaries
of two blossoms united into one fleshy fruit. The more usual
sorts are such as the pine-apple, mulberry, and the fig. These
are, in fact, dense forms of inflorescence, with the fruits or floral
envelopes matted together or coherent with each other ; and all
or some of the parts succulent. The grains of the mulberry
(Fig. 654-656) are not the ovaries of a single flower, like those
G58 657 654
659
656
of the blackberry, which it superficial!}' resembles : the} r belong
to as man}' separate flowers ; and the pulp pertains to the calyx,
not to the pericarp, which is an akene. So that this, like most
multiple fruits, is anthocarpous as well as multiple. Similarly,
the mostl}' indefinite fructiferous masses of Strawberry Elite may
resemble strawberries ; but the pulpy part is the catyx of many
flowers, not the succulent receptacle of one. In the pine-apple,
the flowers are spicate or capitate on a simple axis, which grows
on beyond them into leafy stem ; this when rooted as a cutting
3. Fruit compound (ovaria compound), SYNCARPI ; 4. Collective fruits,
Anthocarpi.
Later, in his "Elements of Botany," Lindley reduced the classes to
two: 1. Simple, Fruits, those proceeding from a single flower; 2. Multiple
fruits, those formed out of several flowers.
FIG. 654. A mulberry, young. 655. One of the fleshy grains at flowering: time, show-
ing it to be a pistillate blossom with fleshy calyx. 656. The same later, with the succu-
lent sepals in transverse section.
FIG. 657. A young fig. 658. Longitudinal section of the same later, but in flowering
time. 659. A small slice, magnified, showing some of the flowers.
ITS KINDS.
303
bears another pine-apple, and so on : the constituent flowers have
through immemorial propagation in this way become sterile and
seedless, and all its parts, along with the bracts and the axis of
the stem, blend in ripening into one flesh}' and juic}' mass. Few
fruits of this class have ever been technically named, at least
with names which have come into use. But the two following
deserve special appellations, although only the latter is familiar
either in ordinary language or in descriptive botany.
584. The Syconium or Hypanthodium, the Fig fruit. (Fig. 657-
659.) This results from a multitude of flowers concealed in a
hollow flower-stalk, if it may be so called, which becomes pulpy
and edible when ripe ; and thus the fruit seems to grow directly
from the axil of a leaf, without being preceded by a blossom.
The minute flowers within, or some of them, ripen their ovaries
into very small akenes, which are commonly taken for seeds.
The fig is to the mulberry what a rose-hip is to a strawberry.
(389, Fig. 406, 407.) It is further explained by a comparison
with a near relative of the Fig-tree, Dorstenia, in which similar
flowers cover the upper surface of a flat peltate disk. This disk
or plate sometimes becomes saucer- shaped b}' an elevation or
incurvation of the margin. A greater degree of this would
render it cup-shaped, or even pitcher-shaped ; from which it is
a short step to the contraction of the mouth down to the small
orifice which is found in the fig.
585. The Strobile or Cone (Fig. 660) is a scaly multiple fruit,
resulting from the ripening of certain sorts of catkin. The name
is applied to the fruit of the Hop, where
the large and thin scales are bracts ;
but it more especially belongs to the
Pine or Fir cone, the peculiar fruit of
Coniferae (507), in which naked seeds
are borne on the upper face of each
fructiferous scale (Fig. 661), or some-
times in their axils.
Such a cone when
spherical, and of
thickened scales
with narrow base, as
that of Cypresses,
has been termed a
GALBULUS, an unnecessaiy name. The galbulus of Juniper is a
FIG. 660. Strobile or Cone of a Pitch Pine, Pinus rigida. 661. Inside view of one
of the scales, showing one of the winged seeds, and the place from which the other, 652,
has been detached.
304 THE FRUIT.
remarkable transformation into a seeming beny ; the few scales
cohering with each other as the}' grow and becoming fleshy at
maturity, completely enclosing a few bony-coated seeds.
586. A Synopsis of the kinds of Fruit, as characterized in
this chapter, is appended. The analysis extends only to simple
fruits. For there are no commonly used special names of
kinds of Aggregate (579), Accessory (580), or Multiple (583)
fruits, except that of Strobile.
SIMPLE FKUITS are
Dry and dehiscent, monocarpellary,
Opening by one (chiefly the ventral) suture, FOLLICLE.
Opening by both sutures, LEGUME.
Or transversely jointed, LOMENT.
Dry and dehiscent, bi-pluri-carpellary, CAPSULE.
When its dehiscence is circumscissile, PYXIS.
When dehiscent by two valves from two parietal placentae, . . SILIQUE.
A short and broad silique, SILICLE.
Dry and bi-pluri-carpellary, splitting into one-seeded carpels, . SCHIZOCARP.
The dimerous schizocarp of Umbelliferae, CREMOCARP.
Each of its halves or carpels, HEMICARP or MERICARP.
The akene-like or nut-like parts into which Schizocarps generally
divide, NUCULES or NUTLETS.
Dry and indehiscent, one-celled, one-two-seeded,
Winged SAMARA.
Wingless, and with the
Thin pericarp consolidated with the seed, CARYOPSIS
Thin pericarp loose and not filled by the seed, UTRICLE.
Thick or hard pericarp free from the seed,
Small, from a one-celled one-two-ovuled ovary, AKENE or ACHENIUM.
Larger, mostly from a two-several-celled and ovuled ovary, . NUT.
Nut borne in a cupule or involucre, GLANS.
Fleshy and indehiscent,
Heterogeneous in texture, having
A stone (putamen) or nutlets within an exterior sarcocarp, . DRUPE.
Papery or cartilaginous carpels in an inferior sarcocarp, . . POME.
A harder or firm rind or exterior, and soft interior,
From an inferior ovary (confined to Gourd Family), . . . PEPO.
From a superior ovary (confined to Orange Family), HESPERIDIUM.
Homogeneous, fleshy throughout, BERRY.
THE SEED. 05
CHAPTER VIII
THE SEED.
oi.v THE SEED is the fertilized ovule (515), with embryo
formed within it. It consists, like the ovule, of a nucleus or
kernel, enclosed by integuments. The seed-coats are those of
the ovule, viz. two, or sometimes only one, in certain plants
none. Occasionally an accessory coat appears after fertiliza-
tion ; and certain appendages may be produced, as outgrowths
from some part of its surface or from its base. The nucleus or
kernel is composed either of the embryo alone, or of a nutritive
deposit in addition. (19-41.) All the parts of a seed are in-
dicated in Fig. 663.
588. The SEED-STALK or PODOSPERM, when there is one, is
the funiculus of the ovule (516), and retains this name. So
also do the CHALAZA, RHAPIIE, and HILUM ; the latter being the
scar left by the separation of the seed from
its funiculus or directly from the placenta.
The foramen of the ovule, now closed, is
the MICROPYLE of the seed.
589. The terms which denote the char-
acter of the ovule, such as orthotropous,
campylotropous, amphitropous, and anatropous, apply equally to
the resulting seed.
590. Seed-Coats. The integuments of the seed answer to the
primine and secundine of the ovule. The main seed-coat is the
exterior integument of the ovule when there is more than one.
Being the most firm coat, and not rarelj' crustaceous in texture,
it takes the name of TESTA, which is equivalent to seed-shell.
It has also been named SPERMODERM (seed-skin), and sometimes
Episperm. The latter name (meaning upon the seed) is best
applied to the pellicle or outer layer, sometimes a thick one,
which the testa of certain seeds forms. The testa is extremely
various in form and texture, is either close and conformed to
FIG. 663. Vertical magnified section of the (anatropous) seed of the American Lin-
den; with the parts indicated, viz. \\w hilum (a); testa (6); tegmen(c); albumen (d);
embryo (*). 664. Vertical section of the orthotropous seed of Helianthemum Cana-
dense, with its funiculus, .
806
THE SEED.
the nucleus, or loose and cellular (as in PjTola-seeds) , or vari-
ously appendaged.
591. The inner coat, called TEGMEN and sometimes ENDO-
PLEUKA, when present is alwa}'s conformed to the nucleus, and
is thin or soft and delicate. Sometimes it is inconspicuous
through cohesion with the nucleus or with the inner surface of
the testa. In ovules of one coat it is necessarily wanting.
592. Appendages or outgrowths of the testa generally have
reference to dissemination. Two characteristic kinds of such
appendages are the wing and the coma,
both pertaining only to the seeds of dehis-
cent fruits and calculated, by rendering
seeds buoyant, to facilitate dispersion by
the wind. The wing of a Pine-seed (Fig.
661, G62) is a part of the carpellary scale
upon which the two ovules grew. In
Trumpet Creeper (Fig. 6G5), an entire
wing surrounds the body of the seed.
In the related Catalpa (Fig. 666), it is
mainly extended from the two ends, and
almost dissolved into a coma, the name
given to the tuft of soft hairs like that
which forms the down at one end of the
seed of Milkweed (Fig. 667), and of
Epilobium, and at both ends in several
Apocynaceae. In the Cotton-plant, very
long and soft hairs, admirably adapted for
iff spinning, thickly cover the whole seed- 057
695 coat. The wing and coma of seeds are
functionally identical with the wing and the pappus of the pericarp
in the samara and the akenes of Composite (563, 564), but
morphologically quite unlike them.
593. There are other (mainly microscopic) structures on some
seed-coats which come usefulty into play in arresting farther
dispersion at a propitious time or place. In many but not all
Polemoniacea? (notably in Collomia), in certain Acanthacese,
such as Ruellia tuberosa (and equally in certain Composite of
the Senecio tribe and in Salvias, &c., among Labiatae, where
this structure is transferred to akenes and nutlets) , the testa is
coated with short hairs, which when wetted burst or otherwise
open and discharge along with mucilage one or more ver} T atten-
FIG. C65. Winged seed of Trumpet Creeper, Tecomaradicans. 6G6. That of Catalpa,
becoming comose: the body divided lengthwise through the embryo.
FIG. 667. Comose seed of Milkweed, Asclepias Cornuti.
ITS COATS AND APPENDAGES.
307
uated long threads (spirides) which were coiled within. These,
protruding in all directions and in immense numbers, form a
limbus of considerable size around the seed, and evidently must
serve a useful end in fixing thsse aual
and light seeds to the soil in time of
rain, or to moist ground, favorable to
germination. In cress and flax-seed,
the abundant mucilage developed when
wetted comes from the gelatination of
epidermal cell-walls, and
subserves a similar use.
594. While the testa in
many seeds is hard and
crustaceous or bony, imitat-
ing the pericarp of a nut, in others (such as Paeonia) it becomes
berry-like (baccate), and in Magnolia, drupaceous. 1 (Fig. 668-
671.) These may also be regarded as adaptations for dissemi-
nation, here by the agency of birds, attracted by bright coloring
and edible pulp.
595. The rhaphe of an anatropous seed (shown in Fig. 681,
685) is sometimes so salient as to form a conspicuous appen-
dage, as in Sarracenia, Fig. 672. Again it may be wholty
1 See article On the Structure of the Ovule and Seed-coats of Magnolia,
in Jour. Linn. Soc. ii. 106, from which the accompanying figures and Fig.
589-597 are reproduced.
FIG. 668. Forming seecl (one eighth of an inch long) of Magnolia Umbrella; the
rhaphe toward the eye. 669. Magnified view of the same divided lengthwise through
the rliaphe; the outer mat, a, beginning to form a hard inner layer, nf. Within and
distinct from this is the inner coat (b\ immediately enclosing the nucleus, c. The oppo-
site side of the testa is thicker on account of the rhaphe, in which d indicates the cord
of spiral ducts.
FIG. 670. A nearly full-grown seed, of the natural size 671. Longitudinal section,
emargeo, snowing the crustaceous or stony inner stratum of the testa well developed:
the parts lettered as in Fig. 669. 672. A transverse section in the same position.
808 THE SEED.
inconspicuous, as in the ripe seed of Magnolia, where it is at
length completely merged and imbedded in the fleshy drupaceous
testa, as shown in Fig. 670-672.
596. Crest-like or other appendages are not uncommon either
on the rhaphe or at the hilum. These are outgrowths produced
during the development of the ovule into the seed. In Sangui-
naria, such a crest develops from the whole length of the rhaphe
(Fig. 673) : in Dicentra, Corydalis (Fig. 674), &c., from some
part of it, mostly from its base next the hilum, or from the
hilum itself, or even from just below it. Such an appendage,
especially when attached to the base of the seed, is named a
STROPHIOLE. A similar and commonly a wart-shaped appendage
in Euphorbia, Ricinus (Fig. 675), &c., is produced by an out-
growth of the external orifice of the ovule, the micropyle of the
seed. This properly takes the name of CARUNCLE. But the
two terms are not always discriminated. By further develop-
ment, either of these may give rise, in certain seeds, to an acces-
sory covering called
597. The Aril or Arillns. This term, rather vaguely employed
by Linnaeus, was first well defined by Gaertner. The true arillus
is an accessor}' seed-covering, more or less incom-
plete, formed between the time of fertilization and
the ripening of the seed, by a growth from the apex
of the funiculus (when there is any) at or just be-
low the hilum, in a manner similar to that in which
the coat or coats of the ovule are formed. That
of Nymphaea (Fig. 676) is a typical example ; only
the arillus is developed from the funiculus at a point distinctly
below its apex : here a ring forms, which grows into a cup, and
this is soon extended into a sac, loosely enclosing the seed, and
open at the top. This is membranaceous ; commonly it is fleshy.
When there is absolutely no. funiculus, the aril may originate
from the placenta, as it does in Podophyllum, in which most of
FIG. 672. Anatropous seed of Sarracenia pnrpurea, with very salient rhaphe. 673.
Same of Sanguinaria or Bloodroot, with rhaphe crested for its whole length. 674. Seed
of Corydalis aurea, with crest or strophiole, attached at or near the hilum. 675. Seed
(suspended) of Ricinus. with its raruncle.
FIG. 676. Seed of White Water-Lily, Nymphsea odorata, in its loose and thin arillus.
AEILLUS, ALBUMEN. 309
the pulp of the beny consists of these flesh}' arils, much com-
pacted. (Fig. 677, G78.)
598. The laciniate aril of the nutmeg (mace) and, it is said,
the bright red and pulpy aril of Euonymus and Celastrus begin
in the manner of a ca-
runcle, and are formed
(mainly if not wholly)
of an outgrowth at or
around the micropyle. So
that, if an orthotropous
seed ever developed an
aril of this sort, it would G--
be seen to begin at the apex of the seed and cover it from above
downward. Planchon, who distinguished this from the true aril,
gave to it the name of ARILLODE (Arillodium) or False Arillus.
599. The Nucleus, or kernel of the seed, consists of the Albu-
men, when this substance is present, and the Embryo.
600. The Albumen, as described in the second chapter (25, &c.) ,
is the name generally employed by systematic botanists for a
store of nutritive matter in the seed outside of the embryo,
whatever its chemical composition. It is not here the name
of a chemical substance (albumen or albumin) , but of a cellular
structure, the cells of which are loaded commonly with starch-
grains (as in the Cerealia) , more or less v mingled with other
matters, or else filled with an encrusting deposit of some equiva-
lent substance, as in the cocoanut, coffee-grain, &c. The cells
in which this deposit is made belong either to the original tissue
of the nucleus, or to a new formation within the embr} T o-sac,
mostly to the latter. (503.)
601. Albumen may be said to belong to all seeds in the grow-
ing stage. In what are called albuminous seeds it persists and
forms either almost the whole kernel, the embryo remaining
minute (as in Fig. 23. 54, 680), or forms a large portion of it (Fig.
13, 17, 19, 21, 48, 663, 664), or, by the growth of the embryo
displacing it, it may in the ripe seed be reduced to a thin stratum
or mere lining to the contiguous seed-coat ; or it may disappear
altogether, as in the seeds of Maple, Almond, Squash, Pea, and
the like, which are therefore said to be exalbuminous. The
difference between albuminous and exalbuminous seeds is that
the maternal nutritive deposit is transferred to the embryo in
FTG. 677. Section of pericarp and placenta of Podophyllum peltatum; the pulp of
the latter mainly of the nature of arillus. investing the seeds. 678. The arillus of ona
seed detached and enlarged, divided lengthwise, showing the seed within.
310 THE SEED.
the former during germination, in the latter during the growth
of the seed.
602. The albumen was named Perisperm by Jussieu, and
Endosperm by Richard (25, note) ; but neither name has in
systematic botany displaced the earlier one of Grew and Gaert-
ner. But both names have recently been brought into use to
distinguish between two kinds of albumen, that formed within
the embiyo-sac, which is specifically termed ENDOSPERM, and that
formed without, which takes the name of PERISPERM. This use
comports with the et} T mology of the two words, the former refer-
ring to a comparatively internal and the latter to an external
portion of the seed or kernel.
603. In most seeds the albumen is endosperm : in Canna it
is all perisperm. In Nymphaea and its allies (except Nelum-
bium, which has none) most of it is perisperm ;
but a thin and condensed layer of endosperm
surrounds the embryo, where with the per-
sistent embryo-sac (or the apex of it) it
forms the fleshy sac in which the embryo is
enclosed. It is the same in the Pepper Family
(Fig. 679) , except that there is a larger quan-
679 tity of endosperm or inner albumen.
604. When the nucleus of a ripe seed is hollow, as in the
cocoanut and mix vomica, the formation of endosperm, which
usually begins next the wall of the embryo-sac, has not proceeded
so as to fill the cavh\y. The embn*o-sac in the cocoanut attains
enormous size, and the cavity is filled by the milky fluid.
605. The texture or consistence of the albumen differs greatly.
It is farinaceous or mealy when, consisting mainly of starch-
grains, it maj- readily be broken down into a powder,
as in wheat, buckwheat, &c. ; oily, when saturated
with a fixed oil, as in popp}'-seed ; fleshy, when
more compact, but readily cut with a knife, as in
the seed of Barberry ; mucilaginous, when soft and
somewhat pulp}', as in Morning Glory and Mallow,
but when dry it becomes fieslry or harder ; corneous,
when of the texture of horn, as in coffee and the
seed of Caulophyllum ; and even bony, as in the vegetable ivory,
the seed of Plrytelephas. It is mostly uniform ; but in the nutmeg,
FIG. 679. Longitudinal magnified section of a seed of Black Pepper; showing the
large episperm, the sinnl! endosperm in the persistent embryo-sac, and in this the
minute embryo.
FIG. 6,0. Longitudinal section of a seed of the so-called Papaw, Asimina triloba,
with ruminated albumen and minute embryo.
THE EMBKYO.
311
in the seeds of Asimina (Fig. 680) and all the Custard- Apple
family, it is marked by transverse lines or divisions (caused by
inflexions or growths of the inner seed-coat) , giving a section
of it either a marbled appearance, or as if it had been slit by
incisions : it is then said to be ruminated.
606. The Embryo, 1 being an initial plantlet or individual of a
new generation, is of course the most important part of the seed.
To its production, protection, and support, all the other parts of
the fruit and flower are subservient.
607. In an embryo of full development, namely, one in which
all the parts are manifest antecedent to germination, these parts
are the Caulicle, otherwise called Radicle, the Cotyledons, and
the Plumule. (20, 30.) The first is the initial
axis or stem, a primaiy internode ; the second
consists of the leaves of the primar} 7 node ; the
third is a beginning of a farther growth which
is to develop more stem and leaves. Such an
embrj'o is usually unaccompanied by albumen,
having in the course of its growth taken into
itself (mostly into the cotyledons) the provision
which in other seeds is mainly accumulated external to it until it
is drawn upon in germination.
1 The word Embryo or Embryon was applied to this body in plants by
Bonnet (Considerations sur les Corps organisees), in 1762, and was introduced
into systematic botany at about the same time (1763) by Adanson : it was
taken up by Gaertner in 1788. Jussieu in the Genera Plantarum (1789) held
to the term Corciilmn (the cor seminis) which came down from Csesalpinus.
Being the germinal part of the seed, the embryo of the plant, like that of
the animal, is in general language often called the Germ.
FIG. 681. Seed of a Violet (anatropous), enlarged; with hilum or scar (a), rhaplie
(ft), and chalaza (r) indicated. (582. Vertical section of the same, showing the straight
embryo in the axis of the mealy albumen.
FIG. 683. Vertical section of the (orthotropous) seed of Buckwheat, showing the
embryo folded round in the mealy albumen.
FIO. 084. Vertical section of the (anatropous) seed of Elodea Virginica, the embryo
completely filling the coats
FIG. 685. Seed of Delphinium tricorne (anatropous), enlarged; the hilum, the
rhaphe, and the chalaza lettered as in Fig. 681. 686. Vertical section of the same with
c, the chalaza, d, the testa, e, the tegmen,/, the albumen, g, the minute embryo near
the hilum (a).
FIG. 687. Embryo of the Pumpkin, with its short radicle and large and flat cotyle
dons, seen flatwise. 688. A vertical section of the same, viewed edgewise.
312 THE SEED.
608. The opposite extreme is an embryo (as in Fig. 68G)
which appears as a mere speck in the albumen, but in which
close microscopical inspection may commonly reveal some differ-
entiation, such as a slight notch at one end (that farthest re-
moved from the microp}ie) of a dicotyledonous embryo, indicating
the future cotyledons. Indeed, in Monotropese, Orobanchacese,
and some other parasitic dicotyledonous plants, and in Orchids
among the monocotyledonous, the embryo is a globular or oblong
particle, with no adumbration of organs whatever antecedent to
germination. There are all grades between the most rudimen-
taiy and the most developed embryos.
609. Under the circumstances of its formation (532), the
radicular end of the embryo is always near to and points towards
the micropyle of the seed, viz. to what was the orifice of the
ovule ; and if the embryo be straight, or merely partakes of the
curvature of the seed, the cotjledons point to the opposite
extremity, that is, to the chalaza.
610. The position of the radicle as respects the hilum varies
with the different kinds of seed. In the orthotropous form, as in
Helianthemum (Fig. 664) and Pepper (Fig. 679), the radicle
necessarily points directly awa3^from the hilum. 1 In the anatro-
pous form, as in Fig. 663, 682, and 684-686, the extremity
of the radicle is brought to the immediate vicinity of the hilum ;
and so it is, although in a different wa}',
in the camp3'lotropous seed (Fig. 689,
690) ; while in the amphitropous the
radicle points away from the hilum later-
ally. As the nature of the ovule and seed
may usually be ascertained by external inspection, so the situation
1 Two technical terms, early introduced by Richard to indicate the direc-
tion of the radicle (caulicle), or rather its relation to the hilum, are
Antitropotis, when the embryo directs its radicle away from the hilum, as
it must in all orthotropous seeds ;
Orthotropous, also homotropous, when directed to the hilum (more strictly to
the micropyle close to the hilum), as in anatropous seeds. These two terms
are still employed by many botanists, although superfluous when the ovule
or seed is stated to be anatropous or orthotropous, &c. And the term
orthotropous, so used, is liable to be confused with orthotropous as applied
to the ovule.
Richard, moreover, termed the embryo amphitropous when curved or coiled,
as in Cliickweed (Fig. 689) and all such campylotropous seeds; and hetero-
ti'ojious when neither radicle nor cotyledons point to the hilum, as occurs
in the semi-anatropous or amphitropous ovule. Many botanists describe
the last by the expression " radicle vague," or, better, " embryo transverse."
FIG. 689. Campylotropous seed of common duckweed, magnified. 690 Section ol
the same, showing the embryo coiled into a ring around the albumen.
THE EMBRYO. 313
of the embiyo within, and of its pails, may often be inferred
without dissection. But the dissection of seeds is not generally
difficult.
611. The direction of the radicle with respect to the pericarp
is also noticed by systematic writers ; who employ the terms
radicle superior or ascending when this points to the apex of the
fruit ; radicle inferior or descending when it points to its base ;
centripetal, when turned toward the axis of the fruit ; centrifugal
(or peritropous] , when turned toward the sides ; and vugue,
when it bears no evident or uniform relation of the kind to the
pericarp.
612. The position of the embr}'o as respects the albumen,
when that is present, is various. Although more commonly in
the axis, it is often excentric, or even external to the albumen,
as in all Grasses and cereal Grains (Fig. 56-61), in PoVygonum,
&c. When external or nearly so, and curved circularly around
the albumen, as in Chickweed (Fig. 690) and Mirabilis (Fig.
17), it is said to be peripheric.
613. The embryo ma} r be very variously folded or coiled in
the seed. The two cotyledons, instead of plane and straight,
may be crumpled ; or they may be simply convolute or rolled up
from one edge, as in Calycanthus (Fig. 691) ; or circinately con-
volute from the apex, as in Bunias ; or else doubled up and thus
biplicately convolute, as in Sugar Maple, Fig. 2. Two modi-
fications are more common, and are of such classificatory impor-
tance in Cruciferae as to need special reference. Namel}', when
cotyledons are
Incumbent (as in Fig. 692, 693), being so folded that the back
of one is laid against the side of the radicle ; and
Accumbent (Fig. 694, 695), when the edges of the pair of
eotj'ledons are longitudinally applied to the radicle. These
differences were first employed in the classification Cruciferse by
FIG. 691. Convolute embryo of Calycanthus, the upper half cut away.
FIG. 692. Seed of a Cruciferous plant (Sisymbriuin), with incumbent cotyledons,
divided. 693. Embryo of the same detached entire.
FIG. 694. Seed of a Cruciferous plant (Barbarea) with accumbent cotyledons
605. The embryo entire.
314 THE SEED.
Robert Brown, and were adopted as primary and tribal characters
by DeCandolle.
614. As to number of cotyledons, the two types of embryo
are the
Monocotyledonous, with a single cot}iedon, i. e. leaves at the
first nodes alternate (39) ; and
Dicotyledonous, with a pair of cotyledons, i. e. leaves of the
first node in the most simple whorl, a pair, in other words, oppo
site (21) ; with its modification of
Polycotyledonous (38), the leaves of the first node in whorls of
three, four, or more. This occurs with constancy in a majority
of Coniferse (Fig. 48, 49), occasionally and abnormally in sundry
ordinary dicotyledonous species.
615. There are several embryos of the cot} T ledonous type in
which one cotj'ledon is smaller than the other, viz. the inner
one when the embiyo is coiled or folded. And in all the species
of Abronia (a genus allied to Mirabilis, Fig. 18) this cotyledon
is wanting, so that the embryo becomes technically monocot}'le-
donous. In another genus, the Dodder (Fig. 78, 79), both
cotyledons are constantly wanting ; and the plumule shows only
minute scales, the homologues of succeeding leaves reduced
almost to nothing.
616. Sometimes the two cotyledons are consolidated into one
body by the coalescence of their contiguous faces ; when they
are said to be conferruminate. This occurs more or less in the
Horsechestnut and Buckeye (Fig. 41, 42), and is striking in
the seed of the Live Oak, Quercus virens.
617. The general morphology of the embryo and its develop-
ment in germination were described at the commencement of this
volume. And so the completion of this account of plant, flower,
fruit, seed, and embryo brings the history round to the starting
point. (12-19, &c.) Having mastered the morphology and
general structure of the higher grade of plants, the pupil may
go on to the morphology and structure of cells (or Vegetable
Anatomy or Histology), and to the study of Cryptogamous
Plants in all their grades.
TAXONOMY. 315
CHAPTER IX.
TAXONOMY.
SECTION I. THE PRINCIPLES OF CLASSIFICATION IN NATURAL
HISTORY.
618. TAXONOMY, from two Greek words which signify arrange-
ment and law, is the study of classification. This is of utmost
importance in Natural Histor}', on account of the vast number
of kinds to be set in order, and of relations (of agreement and
difference) to be noted. Botanical classification, when complete
and correct, will be an epitome of our knowledge of plants.
Arrangement according to kinds, and of special kinds under
the more general, is common to all subjects of stud}'. But the
classification in Biological Natural History, that is in Botany and
Zoolog} T , has a foundation of its own.
619. The peculiarity of plants and animals is that they exist
as individuals, propagating their like from generation to genera-
tion in a series. Of such series of individuals there are very
many kinds, and the kinds have extremely various and unequal
degrees of resemblance. There are various gradations, but not
all gradations of resemblance. Between some, the difference is
so wide that it can be said only that they belong to the same
kingdom ; between others, the resemblance is so close that it
may be questioned whether or not the}' came from common
parents or near ancestors.
620. The recognition of the perennial succession of similar
individuals gives the idea of SPECIES. The recognition of un-
equal degrees of likeness among the species is the foundation
of GENERA, ORDERS, CLASSES, and other groups of species.
621. Individuals are the units of the series which constitute
species. The idea of individuality which we recognize through-
out the animal and vegetable kingdoms is derived from ourselves,
conscious individuals, and from our corporeal structure and
that of the higher brute animals. This structure is a whole,
from which no part can be abstracted without mutilation. Each
individual is an independent organism, of which the component
parts are reciprocally means and ends. Individuality is a main
316 TAXONOMY.
distinction between beings and things ; but, although the tend-
ency to individuation begins with life itself, it is completely
realized only in the higher animals.
622. In plants, as also in some of the lower animals, individu-
ality is merged in commiinity. No plant (except one reduced
to the simplicity of a single cell, of circumscribed growth, and
without organs) is an individual in the sense that a man or a
dog is. (16, 156.) The herb, shrub, and tree are neith-r
indivisible nor of definite limitation. Whether their successive
growths are to remain parts of the previous plant, or to be inde-
pendent plants, depends upon circumstances ; and there is no
known limit to budding propagation.
623. There is, however, a kind of social or corporate indi-
viduality in those animals, or communities (whichever we call
them) of the lower grade which are multiplied by buds or oil-
shoots as well as by ova, and in which the offspring remains, or
may remain, organically connected with the stock. The poly-
pidom or polyparium commonly has a certain limitation and a
definite form ; and certain polyps may become organs with
special functions subordinate to the common weal. This is
more largely true in the vegetable kingdom. So that for de-
scriptive purposes, and in a just although somewhat loose sense,
the herb, shrub, or tree is taken as an individual. But only
while it forms one connected body. Offshoots when separately
established are equally individuals in this sense.
624. What it is in plants which philosophically answers to the
individual in the higher animals is another question, to which
various answers have been given. 1 Some insist that the whole
vegetative product of one seed makes one individual, whether
connected or separated (as may happen) into a million of plants.
But a common and less strained view restricts the individual
to such product only while organically united. Others (of
which Thouars at the beginning and Braun at the middle of the
present centuiy are leading examples) take each axis or shoot
with its foliage to represent the individual, of which the leaves
and their homologues are organs, the branches being usually
implanted upon the parent axis as this is implanted in the soil,
but also equally capable of producing roots by which they may
make their own connection with the soil. Still others, on pre-
1 For the history of opinion upon and a full presentation of this topic,
see Alexander Braun's Memoir (originally published in the Abhandl. Akad.
Wissenschaften zu Berlin, 1853), Das Individuum der Pflanze, &c., and a
translation by C. F. Stone in Amer. Jour. Sci. ser. 2, xix. xx. 1855.
THE PRINCIPLES OF CLASSIFICATION. 317
cisely similar grounds, carry the analysis a step farther, and
regard each phytomer (16) as the individual. Finally, some, in
view of their potentially independent life, take the cells, or units
of anatomical structure, to be the true individuals ; and this
with sufficient reason as regards the simplest cryptogamous
plants. Upon the view here adopted, that plants do not rise
high enough in the scale of being to reach true individuality,
the question is not whether it is the cell, the phytomer, the shoot,
the tree, or the whole vegetative product of a seed which answers
to the animal individual, but only which is most analogous to it.
In our view, its analogue is the cell in the lowest grades of vege-
table life, the phytomer in the higher. 1 But, in botanical de-
scription and classification, by the individual is meant the herb,
shrub, or tree, unless otherwise specified .
625. Species in biological natural histoiy is a chain or series
of organisms of which the links or component individuals are
parent and offspring. Objectively, a species is the totality oi.-
beings which have come from one stock, in virtue of that moU
general fact that likeness is transmitted from parent to progeny.
Among the many clelinitions, that of A. L. Jussieu is one of the
briefest and best, since it expresses the fundamental conception
of a species, i. e. the perennial succession of similar individuals
perpetuated by generation.
626. The two elements of species are : 1 , community of origin ;
and, 2, similartty of the component individuals. But the degree
of similarity is variable, and the fact of genetic relationship can
seldom be established by observation or historical evidence. It
is from the likeness that the naturalist ordinarily decides that
such and such individuals belong to one species. Still the like-
ness is a consequence of the genetic relationship ; so that the
latter is the real foundation of species.
1 For just as successive branches are repetitions and progeny of the
parent brancli or stem, the phytomers of the branch are repetitions and
progeny each of the preceding one, so forming a series of vegetative
generations ; and the whole tree might almost as well represent the individ-
ual as one of its branches. The phytomer, as well as the branch, is capable
of completing itself by producing roots, but is itself indivisible except by
mutilation. Least tenable of all is the conception that the whole product
of a seed may be taken to represent the vegetable individual. For then
individuals increased by buds and division are wholly unlimited both in ex-
tent and in duration, so far as observation can show, and a multitudinous
race, not only of the present and past, but perhaps in perpetuity, may con-
sist of a single individual. There are, indeed, theoretical reasons for infer-
ring that a bud-propagated race may not last so long as a seed-propagated
species ; but there is no proof of it. See Darwiniana, Art. xii.
318 TAXONOMY.
627. No two individuals are exactly alike ; and offspring of
the same stock may differ (or in their progeny may come to differ)
strikingly in some particulars. So two or more forms which
would have been regarded as wholly distinct are sometimes
proved to be of one species by evidence of their common origin,
or more commonly are inferred to be so from the observation of
a series of intermediate forms which bridge over the differences.
Only observation can inform us how much difference is compat-
ible with a common origin. The general result of observation
is that plants and animals breed true from generation to genera-
tion within certain somewhat indeterminate limits of variation ;
that those individuals which resemble each other within such
limits interbreed freely, while those with wider differences do
not. Hence, on the one hand, the naturalist recognizes Varieties
or differences within the species, and on the other Genera and
other superior associations, indicative of remoter relationship of
the species themselves.
G2H. Varieties are forms of species marked by characters of
less fixity or importance than are the species themselves. The}'
may be of all grades of difference from the slightest to the most
notable : they abound in free nature, but assume particular
importance under domestication and cultivation ; under which
variations are most prone to originate, and desirable ones are
preserved, led on to further development, and relatively fixed.
G29. If two seeds from the same pod are sown in different
soils, and submitted to different conditions as respects heat, light,
and moisture, the plants that spring from them will show marks
of this different treatment in their appearance. Such differences
are continually arising in the natural course of things, and to
produce and increase them artificially is one of the objects of
cultivation. Striking as they often are (especially in annuals
and biennials) , they are of small scientific consequence. When
spontaneous they are transient, the plant either outlasting the
modifying cause or else succumbing to its continued and graver
operation. But, in the more marked varieties which alone de-
serve the name, the cause is occult and constitutional ; the
deviation occurs we know not why, and continues throughout
the existence and growth of the herb, shrub, or tree, and con-
sequently through all that proceeds from it by propagation from
buds, as by offsets, ^ers, cuttings, grafts, &c.
G30. Some varieties of cultivation originate in comparatively
slight deviations from the type, and are led on to greater differ-
ences by strict selection of the most marked individuals to
breed from. Most appear as it were full-fledged, except as to
THE PKINCIPLES OF CLASSIFICATION. 319
luxuriance or development, more or less under the control of
conditions, their origin being wholly unaccountable. They arise
in the seed-bed, or sometimes from buds, which as the gardeners
say " sport" l That is, some seedlings, or some shoots, are
unlike the rest in certain particulars. 2
631. Most varieties originate in the seed, and therefore the
foundation for them, whatever it may be, is laid in sexual repro-
duction. But Bud-variation, or the u sporting" of certain buds
into characters in branch, flower, or fruit unlike those of the
stock, is known in a good number of plants. 3 It might also
o^cur in corals, hydras, and other compound animals propagated
by budding. Once originated, these varieties mostly persist,
like seedling varieties, through all the generations of budding
growth, but are not transmitted to the seed.
632. Upon the general principle that progeny inherits or tends
to inherit the whole character of the parent, all varieties must
have a tendency to be reproduced by seed. But the inheritance
of the new features of the immediate parent will commonly be
overborne by atavism, i. e. the tendency to inherit from grand-
parents, great-grand-parents, &c. Atavism, acting through a long
line of ancestry, is generally more powerful than the heredity
of a single generation. But when the offspring does inherit the
peculiarities of the immediate parent, or a part of them, its off-
spring has a redoubled tendency to do the same, and the next
generation still more ; for the tendency to be like parent, grand-
parent, and great-grand-parent now all conspire to this result
and overpower the influence of remoter ancestry. Close-breed-
ing (398) is requisite to this result. In the natural wild state,
varieties many and conspicuous as they often are must be
much repressed by the prevalent cross-fertilization which takes
place among the individuals of almost all species. Cultivators and
breeders in fixing varieties are careful to secure close breeding
as far as this is possible. This has fixed the particular sorts of
Indian Corn, Rye, Cabbage, Lettuce, Radishes, Peas, &c., and
1 Both the technical English term. Sport, and its Latin equivalent, Lusus,
are sometimes used for bud-variation only, yet as commonly for seedling
variation also.
' 2 Darwin assumes that variation is of itself indefinite or vague, tending
in no particular direction, but that direction is wholly given by the elimina-
tion in the struggle for life of all but the fittest for the conditions. But
what we observe in the seed-bed does not suggest this view. Naageli, Braun,
and myself incline to the opinion that each plant has an inherent tendency
to variation in certain general directions.
3 A list of known bud-varieties is given in Darwin's Variation of Animals
and Plants under Domestication, Chapter xi.
320 TAXONOMY.
indeed of nearly all our varieties of cultivated annual and biennial
esculent plants, as well as of several perennials, many of which
have been fixed through centuries of domestication, while others
are of recent establishment. What is now taking place with
the Peach in this country ma}- convince us that heritable varieties
ma}' be developed in trees as well as in herbs, and in the same
manner ; and that the reason why most races are annuals or
biennials is because these can be perpetuated in no other way,
arid because the desired result is obtainable in fewer years than
in shrubs or trees. Varieties of this fixity of character are called
633. Races (Lat. Proles). A race, in this technical sense of
the term, is a variety which is perpetuated with considerable
certainty by sexual propagation. This distinction of varieties
pertains chiefly to botany. In the animal kingdom all permanent
varieties must be races. So are all indigenous varieties of
plants. 1 In most of these, the position of species and variety is
more or less arbitrary or accidental, and capable of interchange.
What is called the species may be only a commoner or better-
known form, or the one first recognized and named by botanists ;
whence the other forms as they come to be recognized are made
to rank in the books as varieties. Instead of one varying from
the other, all the forms have probably varied ages ago from a
common t}*pe.
634. These varieties of the highest order and most marked
characteristics, being perpetuable by seed, have the principal
attributes of species. They are a kind of subordinate derivative
species. Hence they are sometimes called Subspecies. We
judge them not to be so many species, either because in the case
of cultivated races we know something of their origin or histor}',
and more of the grave changes which long domestication may
bring to pass ; or because the forms, however stable, differ
among themselves less than recognized species generally do ; or
because very striking differences in the extremes are connected
by intermediate forms. And our conclusions, it must be under-
stood, "are not facts, but judgments, and largely fallible judg-
ments." 2 For while some varieties appear strikingly different,
some species are very much alike. 3
1 The Horseradish and a few other plants of spontaneous growth, which
through long dependence on bud-propagation seem to have lost the power
of setting seed, can hardly be called varieties.
2 Darwiniana, 35.
8 Wherefore, since we hardly need the term race in the restricted sense
of seed-propagated variety, it is sometimes convenient to use it in the man-
ner proposed by Bentham (Anniversary Address to the Linnean Society,
THE PRINCIPLES OF CLASSIFICATION. 321
635. One distinction between varieties and species is note-
worthy and important, even if it may not serve as a criterion.
The individuals of different varieties in plants interbreed as freely
as do those of the same variety and are equally prolific. Their
union produces
636. Cross-breeds. 1 In nature, cross-breeding doubtless re-
presses variation or prevents the segregation of varieties into
what would be ranked as species. In cultivation and domesti-
cation, it is turned to important account in producing intermediate
new varieties (cross-breeds) variously combining the different
excellencies of two parent individuals or two varieties. Thus
the great number of forms produced by variation (especially as
to flowers and fruits) have been further diversified, and selected
forms improved for special uses by judicious combination.
637. In general, the individuals of distinct species do not
interbreed, although many are capable of it. There is great
diversity in this regard among plants, some (such as Willows,
Verbascums, and Verbenas) interbreeding freely and reciprocally ;
some interbreeding in one direction, but not reciprocally ; others,
even when very similar, refusing to unite. But, on the whole,
there seems to be few nearly related species in which the pollen
of the one cannot be made to act upon the ovules of the other
by persistent and proper management. Such crossing is an
important resource in horticulture. Crossing of species, when
successful, produces
638. Hybrids. In these, the characteristics of the two species
are combined, sometimes in equal proportions, sometimes with
great preponderance of one or the other parent ; and there is
often a difference in the result in reciprocal fertilizations. Hy-
brids do not play a very prominent part in nature, apart from
cultivation, although the limits of some species may be obscured
by them, possibly of more than is generally supposed. In the
animal kingdom, all the most familiar hybrids are sterile : in the
vegetable kingdom, a majority may have a certain but very low
degree of fertility ; but this is also the case in many unions
May, 1869, 5) as the common designation of any group or collection of indi-
viduals whose characters are continued through successive generations,
whether it be permanent variety, subspecies, species, or group consisting
of very similar species, the term not implying any decision of this question.
If this use of the term race prevails, Subspecies will probably take its place
as the designation of the highest grade of variety. The objection to this is
that the subspecific and specific names would be more liable to be confused.
1 Half-breed is a common equivalent term in the animal kingdom : Latin,
Mistus or Mixtas ; French, Metis.
21
322 TAXONOMY.
within the species, and especially in the application of the pollen
to the stigma of the same blossom. Commonly the sterilit}* of
hybrids is owing to the impotence of the stamens, which perfect
no pollen ; and most such hybrids may be fertilized by the pollen
of the one or the other parent. Then the offspring either in
the first or second generation reverts to the fertilizing species.
Moreover, certain hybrids, such as those of Datura, which are
fully fertile per se, divide in the offspring, partly in the first gen-
eration, and completely in two or three succeeding generations,
into the two component species, even when close-fertilized. 1 (In
part this may come from adventive embryo-formation, 533.)
639. There appears, therefore, to be a real ground in nature
for species, notwithstanding the difficulty and even impossibility
in many cases of defining and limiting them.
640. Species is taken as the unit in zoological and botanical
classification. Important as varieties are in some respects,
especially under domestication and cultivation, they figure in
scientific arrangement only as fractions of species. Species
are the true subjects of classification. The aim of sj^stematic
natural history is to express their relationship to each other.
641. The whole ground in nature for the classification of spe-
cks is the obvious fact that species resemble or differ from each
other unequally and in extremely various degrees. If this were
not so, if related species differed one from another by a constant
quantity, so that, when arranged according to their resemblances,
the first differed from the second about as much as the second
from the third, and the third from the fourth, and so on, or if
the species blended as do the colors of the rainbow, then, with
all the diversitj" in the vegetable kingdom there actually is, there
could be no natural foundation for their classification. The mul-
titude of species would render it necessary to classify them, but
the classification would be wholly artificial and arbitrary. The
actual constitution of the vegetable kingdom, however, as ap-
pears from observation, is that some species resemble each other
veiy closely indeed, others differ as widely as possible, and be-
tween these the most numerous and the most various grades of
1 According to Naudin in Comptcs Rendus, xlix. 1859, & Iv. 1862. See
also Naudin's memoir on hybridity in plants in Ann. Sci. Nat. ser. 4, xix.
1863, pp. 180-203, & in Mem. A cad. S^-i. . . . For the literature on vegetable
hybrids, see Kcelreuter, Nachricht, &c., 1761, and Appendices, 1763-1766;
Herbert, on Amaryllidaceae, 1837 ; C. F. Gasrtner, Versuche und Beobachtun-
gen ueber die Bastarderzeugung in Pflanzenreich, 1849; Wichura, Die
Bastardbefruchtung im Pflanzenreich, erlautertert an den Bastarden der
Weiden ; and the mnmoir of Naudin referred to.
THE PRINCIPLES OF CLASSIFICATION. 3^3
resemblance or difference are presented, but alwaj's with a mani-
fest tendency to compose groups or associations of resembling
species, groups the more numerous and apparently the less
definite in proportion to the number and the nearness of the
points of resemblance. These various associations the naturalist
endeavors to express, as far as is necessary or practicable, l>y a
series of generalizations, the lower or particular included in the
highar or more comprehensive. All kinds of differences are
taken into account, but only the most constant and definite ones
are relied on for characters, i.'e. distinguishing marks. Linnaeus
and the naturalists of his day used names for only three grades
of association, or groups superior to species, viz. the Genus, the
Order, and the Class ; and these are still the principal members
of classification.
642. Genera (plural of Genus) are the more particular or
special groups of related species. They are groups of species
which are much alike in all or most respects, which are con-
structed, so to say, upon the same particular model, with only
circumstantial differences in the details. They are not neces-
sarily nor generally the lowest definable groups of species, but
are the lowest most clearly definable groups which the botanist
recognizes and accounts worthy to bear the generic name ; for
the name of the genus with that of the species added to it is the
scientific appellation of the plant or animal. Constituted as the
vegetable and animal kingdoms are, the recognition of genera, or
groups of kindred species, is as natural an operation of the mind
as is the conception of species from the association of like indi-
viduals. This is because many genera are so strongly marked,
at least so far as ordinary observation extends. Every one
knows the Rose genus, composed of the various species of Roses
and Sweetbriers ; the Bramble genus, comprising Raspberries,
Blackberries, &c., is popularly distinguished to a certain extent ;
the Oak genus is distinguished from the Chestnut and the Beech
genus ; each is a group of species whose mutual resemblance is
greater than that of any one of them to an} r other plants. The
number of species in such a group is immaterial, and in fact is
very diverse. A genus may be represented by a single known
species, when its peculiarities are equivalent in degree to those
which characterize other genera. This case often occurs ; al-
though, if this were universally so, genus and species would be
equivalent terms. If only one species of Oak were known, the
Oak genus would have been as explicitly discerned as it is now
that the species amount to three hundred ; and better defined,
for now there are forms quite intermediate between Oak and
324
TAXONOMY'.
Chestnut. Familiar illustrations of genera in the animal king-
dom are furnished by the Cat kind, to which belong the domestic
Cat, the Catamount, the Panther, the Lion, the Tiger, the Leop-
ard, &c. ; and by the Dog kind, which includes with the Dog
the different species of Foxes and Wolves, the Jackal, &c. The
languages of the most barbarous as well as of civilized people
ever}' where show that the}" have recognized such groups. Natu-
ralists merely give to them a greater degree of precision, and
indicate what the points of agreement are.
643. If most genera were as conspicuously marked as those
from which these illustrations are taken, genus would be as defi-
nitely grounded in nature as species. But popularly recognized
genera, rightty based, are comparatively few. Popular nomen-
clature, embodying the common ideas of people, merely shows
that generic groups are recognizable in a considerable number
of cases, but not that the whole vegetable or the whole animal
kingdom is divisible into a definite number of such groups of
equally or somewhat equally related species. The naturalist
discerns the ground of genera in characters which the casual and
ordinary observer overlooks ; and, taking the idea of genera
from the numerous well-marked instances as the norm, applies
it as well as possible to the less obvious or less natural cases, and
groups all known species under genera. Resemblances among
the species when rightly grouped into genera, though real, are
often so unequal in degree, that certain species may be about as
nearly related to neighboring genera. So that the recognition of
genera even more than of species is a matter of judgment, and
even of conventional agreement as to how and where a certain
genus shall be limited, and what particular association of species
shall hold the position of genus. All the species of a genus must
accord in every important structure ; but extended observation
only can settle the question as to what are important and what
are incidental characters. For example, the pinnatifid or sinu-
ate leaf might have been thought as essential to the Oak genus
as the acorn-cup ; but many Oaks are now known with entire
leaves, resembling those of Willow or Laurel. An open acorn-
cup beset with imbricated scales is a character common to all
European and American Oaks ; but in numerous Asiatic species
the cup bears concentric or spiral lamellae instead, and in others
the cup takes the form of a naked and closed sac. Maples have
palmately-veined and lobed leaves ; but one species has undi-
vided and pinnately-veined leaves. The Apple and the Pear
under one view are of the same genus, under another they rep-
resent different genera.
THE PRINCIPLES OF CLASSIFICATION. 325
644. The genus must be based on close relationship of species,
but not necessarily on the closest. Raspberries differ from
Blackberries, but must be ranked in the same genus ; and so of
Plums and Cherries. For the groups which are to bear the
generic name must be as distinct and definite as possible.
645. Orders are to genera what genera are to species. They
are groups of a higher rank and wider comprehension, expressive
of more general resemblances, or, in other language, of remoter
relationship. As all species must be ranked in genera, so all
genera must be ranked in orders. FAMILY in botany is synony-
mous with order : at least natural orders and families (however
distinguished in zoology) have always in botany been inter-
changeable terms, and will probably so continue. 1
646. As examples of orders in the vegetable kingdom take
the Oak family, composed of Oaks, Chestnuts, Beeches, &c. ;
the Pine family, of Pines, Spruces, Larches, Cedars, Araucaria,
Cypresses, and their allies ; the Rose family, in which Brambles,
Strawberries, Plums and Cherries, Apples and Pears are asso-
ciated with the Rose in one somewhat multifarious order.
647. Classes are to orders what these are to genera. They
express still more comprehensive relations of species ; each class
embracing all those species which are framed upon the same
broad plan of structure, however differently that plan may be
carried out in particulars.
648. Kingdom must be added, to represent the highest gener-
alization. All subjects of biological classification belong either
to the vegetable or animal kingdom. Mineralogy, Chemistry,
&c., ma}' use the same terms (genus, species, &c.) in an analo-
gous way ; but the classification of substances rests on other
foundations than that of beings.
649. The sequence of groups, rising from particular to univer-
sal, is Species, Genus, Order, Class, Kingdom ; or, in descending
from the universal to the particular,
KINGDOM,
CLASS,
ORDER,
GENUS,
SPECIES.
1 Order is the older term, and that which associates best with the technical
Latin names. Family is a happy term, which associates itself well with
English names. But its use is attended with this incongruity, that the tribe
(653) in natural history classification is subordinate to the family. In
zoology, order is distinguished from family as the next higher grade.
326 TAXONOMY.
650. This is the common framework of natural history classi-
fication. All plants and all animals belong to some species ;
every species to some genus ; every genus to some order or
family ; every order to some class ; every class to one or the
other kingdom. 1 But this framework, although all that is re-
quisite in some parts of natural history, does not express all the
observable gradations of relationship among species. And even
gradations below species have sometimes to be classified. The
series is capable of extension ; and extension is often requisite
on account of the large number of objects to be arranged, and
the various degrees of relationship which may come into view.
651. This is effected by the intercalation of intermediate
grades, to be introduced into the system only when there is
occasion for them. And in botan}^ one or more grades superior
to the classes are needful ; for first and foremost is the great
division of all plants into a higher and a lower SERIES 2 (or sub-
kingdom) , the Phaenogamous and the Cryptogamous.
652. The grades intercalated into the long-established sequence
of Class, Order, Genus, and Species, with new names, are mainly
two, Tribe and Cohort.
653. Tribe has been for a generation or two thoroughly estab-
lished in both kingdoms, as a grade inferior to order and supe-
rior to genus. In botanical classification, much use is made of
this grade, genera being grouped into tribes.
654. Cohort (Lat. Cohors) is of more recent introduction, at
least in Botany, but is becoming established for a grade next
above that of order. Orders are grouped into cohorts. Lindley
hit upon a good English name for this grade, that of Alliance.
But this word has no available Latin equivalent ; while cohort
takes equally well a Latin or an English form.
655. Finally, each grade is capable of being doubled by the
recognition of one like it and immediately subordinate to it, and
with designation directly expressive of the subordination. For
1 Not recognizing Haeckel's third kingdom of Protista, consisting of those
lowest forms of being from which the animal and vegetable kingdoms
emerge.
2 Answering to the French Embranchement in zoology. For this it is pro-
posed to use the word Division (Divisio) : see Laws of Botanical Nomencla-
ture adopted by the International Botanical Congress held at Paris in
August, 1867 ; together witli a Historical Introduction and a Commentary, by
Alph. DeCandolle, English translation, London, 1868 ; the original French
edition, Paris, 1867. Perhaps no better name can be found ; but the elder
DeCandolle brought Dirisio into common use for a grade subordinate to
tribe. Endlicher employed the term Regio. We have used Series, and
much prefer it.
THE PRINCIPLES OF CLASSIFICATION. 827
example, if Dicot3 r ledones and Monocot3iedones be the two
classes of Phaenogamia, the former (and only the former) is
divided upon very important characters into two branches, of far
higher rank than the cohorts, viz. the Angiospermae and the
Gymnospermae, which take the name of SUBCLASSES. Orders,
especially the more comprehensive ones, often comprise two or
more groups so distinct that it may fairly be a question whether
they are not of ordinal rank : such take the name of SUBORDERS.
Tribes in like manner may comprise groups of similar relative
value : these are SUBTRIBES. Genera ma}* comprise sections of
species which might almost as well rank as genera themselves :
to mark their importance and pretension (which may come to
be allowed) , they are termed SUBGENERA. Finally, forms which
are ranked as varieties, but which may establish a claim to be
distinct species, are sometimes termed SUBSPECIES. Even what
we regard as a variety may comprise more or less divergent
forms, to be distinguished as SUBVARIETIES.
656. Some of the larger and most diversified orders, tribes,
genera, or species may require all these analytical appliances,
and even more, for their complete elucidation ; while others,
comparatively homogeneous, offer no ground for them. But
when these grades, or some of them, come into use, they are
always in the following sequence :
KINGDOM,
SERIES or Division, or Sub-kingdom,
CLASS,
Subclass,
Cohort,
ORDER or Family,
Suborder,
TRIBE,
Sub tribe,
GENUS,
Subgenus,
Section,
Subsection,
SPECIES,
Subspecies or Race,
Variety,
Subvariety.
657. Nature and Meaning of Affinity. These grades, the higher
including the lower, denote degrees of likeness or difference.
Plants belonging respectively to the two great series or primary
divisions may accord only in the most general respects, in that
which makes them plants rather than animals. Plants of the
same variety are generally as much alike as if they were of the
328 TAXONOMY.
same immediate parentage. All plants of the same species are
so much alike that the}- are inferred to have descended from
a common stock, and their differences, however grave, are sup-
posed to have arisen from subsequent variation, and the more
marked differences to have become fixed through heredity. This
is included in the idea of species. Descent from a common
origin explains the likeness, and is the only explanation of it.
658. But what is the explanation of the likeness between the
species themselves? As respects nearly related species, the answer
is clear. Except for practical purposes and in an arbitraiy
wa}', no certain and unfailing distinction can be drawn between
varieties of the highest grade and species of closest resemblance.
It cannot reasonably be doubted that they are of similar origi-
nation. Then there are all gradations between very closely and
less closety related species of the same genus of plants.
659. The Theory of Descent, that is, of the diversification of the
species of a genus through variation in the lapse of time, affords
the only natural explanation of their likeness which has }'et been
conceived. The alternative supposition, that all the existing
species and forms were originally created as they are, and have
come down essentially unchanged from the beginning, offers no
explanation of the likeness, and even assumes that there is no
scientific explanation of it. The Irypothesis that the species of
a genus have become what the} T are by diversification through
variation is a very old one in botany, and has from time to time
been put forward. But until recently it has had little influence
upon the science?, because no clear idea had been formed of any
natural process which might lead to such result. Doubtless, if
variation, such as botanists have to recognize within the species,
be assumed as equally or even more operative through long ante-
rior periods, this would account for the diversification of an
original species of a genus into several or many forms as differ-
ent as those which we recognize as species. But this would not
account for the limitation of species, which is the usual (but
not universal) characteristic, and is an essential part of the
idea of species. Just this is accounted for by
660. Natural Selection. This now familiar term, proposed by
Charles Darwin, was suggested by the operations of breeders in
the development and fixation of races for man's use or fane}' ;
in animals by breeding from selected parents, and selecting for
breeding in each generation those individuals only in which the
desired points are apparent and predominant ; in the seed-bed
by rigidly destroying all plants which do not show some desirable
variation, breeding in and in from these, with strict selection of
THE PRINCIPLES OF CLASSIFICATION. 329
the most variant form in the particular line or lines, until it be-
comes fixed by heredity and as different from the primal stock
as the conditions of the case allow. In nature, the analogous
selection, through innumerable generations, of the exceedingly
small percentage of individuals (as ova or seeds) which ordi-
narily are to survive and propagate, is made by competition for
food or room, the attacks of animals, the vicissitudes of climate,
and in fine by all the manifold conditions to which they are
exposed. In the Struggle for Life to which they are thus inevi-
tably exposed, only the individuals best adapted to the circum-
stances can survive to maturity and propagate their like. This
Survival of the Fittest, metaphorically expressed by the phrase
natural selection, is in fact the destruction of all weaker com-
petitors, or of all which, however they might be favored by other
conditions, are not the most favored under the actual circum-
stances. But seedlings varying, some in one direction, some in
another, are thereb}' adapted to different conditions, some to one
kind of soil or exposure, some to another, thus lessening the com-
petition between the two most divergent forms, and favoring their
preservation and farther separation, while the intermediate forms
perish. Thus an ancestral type would become diversified into
races and species. Earlier variation under terrestrial changes
and vicissitudes, prolonged and various in geological times since
the appearance of the main types of vegetation, and the attendant
extinctions, are held to account for genera, tribes, orders, &c.,
and to explain their actual affinities. Affinity under this view
is consanguinity ; and classification, so far as it is natural, ex-
presses real relationship. Classes, Orders, Tribes, &c., are the
earlier or main and successful branches of the genealogical tree,
genera are later branches, species the latest definitely developed
ramifications, varieties the developing buds. 1
6G1. Except as to those changes in size, luxuriance, or depau-
peration and the like, in which plants, especially seedlings,
respond promptly to external influences, as to heat or cold,
1 For the inception of this theory of descent in the form which has within
the last twenty years profoundly affected natural history, and developed a
copious literature, see a short paper On the Variation of Organic Beings in
a State of Nature ; On the Natural Means of Selection ; and On the Com-
parison of Domestic Races and True Species, by Charles Darwin, also On
the Tendency of Varieties to depart indefinitely from the Original Type, by
Alfred Russell Wallace, both read to the Linnean Society, July 1, 1858, and
published in its Journal of the Proceedings, iii. (Zoology) 45-62. For the
development of the doct: ine, soe Darwin's " Origin of Species by Means of
Natural Selection," "The Variation of Animals and Plants under Domes-
tication," and various other works; Wallace's "Geographical Distribution
of Animals," &c. For some expositions, see Gray's " Darwiniana."
330 TAXONOMY.
moisture or diyness (which are transient and comparative!}*
unimportant), variation, or the unlikeness of progen}* to parent,
is occult and inexplicable. If sometimes called out by the
external conditions, it is by way of internal response to them.
In Darwin's conception, variation of itself does not tend in any
one particular direction : he appears to attribute all adaptation
to the sorting which results from the struggle for existence and
the survival of the fittest. We have supposed, and Nsegeli takes
a similar view, 1 that each plant has an internal tendency or pre-
disposition to vary in some directions rather than others ; from
which, under natural selection, the actual differentiations and
adaptations have proceeded. Under this assumption, and taken
as a working hypothesis, the doctrine of the derivation of species
serves well for the co-ordination of all the facts in botany, and
aifords a probable and reasonable answer to a long series of
questions which without it are totally unanswerable. It is sup^
ported by vegetable palaeontology, which assures us that the
plants of the later geological periods are the ancestors of the
actual flora of the world. In accordance with it we may explain,
in a good degree, the present distribution of species and other
groups over the world. It rationally connects the order of the
appearance of vegetable t}"pes in time with the grades of differ-
entiation and complexity, both proceeding from the simpler, or
lower and more general, to the higher and more differentiated
or special ; it explains by inheritance the existence of function-
less parts ; throws light upon the anomalies of parasitic plants in
their various gradations, upon the assumption of the most various
functions by morphological ly identical organs, and indeed illumi-
nates the whole field of morphology with which this volume has
been occupied. It follows that species are not " simple curiosities
of nature," to be catalogued and described merely, but that they
have a history, the records of which are impressed upon their
structure as well as traceable in their geographical and palaeon-
tological distribution. This view, moreover, explains the re-
markable fact that the characters in which the affinities of
plants are mainly discerned (and which therefore serve best
for orders, tribes, and other principal groups) are commonly such
as are evidently of small if any importance to the plant's well-
being, and that they run like threads through a series of species
of the greatest diversity in habit, mode of life, and particular
adaptations to conditions. 2
1 Entstehung und Begriff der naturhistorischen Art. Zweite Auflage, 1805.
2 This is a corollary of natural selection, which can take effect only
upon useful characters, i. e. upon structures which play some active part
BOTANICAL CLASSIFICATION. 831
662. The fixity of species under this view is not absolute and
universal, but relative. Not, however, that speciiic changes are
necessitated in virtue of any fixed or all-controlling natural law.
Some of the lowest forms have existed essentially unchanged
through immense geological periods down to the present time ;
some species even of trees are apparently unchanged in the lapse
of time and change of conditions between the later tertiary period
and our own day, during which most others have undergone
specific modification. Such modifications are too slow to effect
in any wise the stability and practical application of botanical
classification.
SECTION II. BOTANICAL CLASSIFICATION.
633. Natural and Artificial Classifications may be distinguished.
A natural classification in botany aims to arrange all known plants
into groups in a series of grades according to their resemblances,
and their degrees of resemblance, in all respects, so that each
species, genus, tribe, order, &c., shall stand next to those which
it most resembles in all respects, or rather in the whole plan of
structure. For two plants may be very much alike in external
appearance, yet very different in their principal structure. Arti-
ficial classifications single out one or more points of resemblance
or difference and arrange by those, without reference to other
considerations, convenience and facility being the controlling
principles. The alphabetical arrangement of words in a dic-
tionary, and the sexual system in botany by Linmcus (or rather
a part of it) , in which plants are arranged in classes upon the
number of their stamens, and in orders upon the number of
pistils, are examples of artificial classification. The arrange-
ment of the words of a language under their roots, and with the
derivative under the more primitive forms, would answer to a
natural classification.
in the life of the plant, and which therefore undergo modification under
changing conditions. Unessential structures accordingly are left unaltered
or are only incidentally modified. And so these biologically unessential
points of structure, persisting through all adaptive changes, are the clews
to relationship. Thus, Rubiaceae are known by insignificant stipules, Ano-
naceae by ruminated albumen, Tlhamnacea? by a valvate calyx and stamens
before the petals, &c. Paradoxical as it may seem, it is not although, but
because, they are of small biological importance that they are of high clas-
sificatory (i. e. of genealogical) value.
On considerations like these, characters are divided into adaptive or bio-
logical on the one hand, and genealogical or genetic on the other. The saga-
cious naturalist seizes upon the latter for orders and the like ; while the
former are prominent in genera, &c.
332 TAXONOMY.
664. No artificial classification of plants could fail to be
natural in some portions and some respects ; because plants
which agree in any point of structure likely to be used for the
purpose will commonly agree in other and perhaps more impor-
tant characters. On the other hand, no natural classification can
dispense with artificial helps ; nor can it express in lineal order,
or in any other way, all the various relationships of plants, even
if these were full}' determined and rightl} r subordinated. Natu-
ralists now endeavor to make classification as natural as possible ;
that is, to base it in every grade upon real relationships. What
real relationships are, and how to express them in a general
system and throughout its parts, has been the task of the leaders
in botany from the beginning of the science until now ; and the
work is b} r no means completed.
665. Linnaeus was perhaps the first botanist to distinguish
clearly between a natural and an artificial classification. He
labored ineffectually upon a natural classification of the genera
of plants into orders ; and he devised an effective artificial classi-
fication, which became so popular that it practically superseded
all others for more than half a century, and has left a permanent
impression upon the science. The last generation of botanists
who were trained under it has not quite passed away.
666. Ante-Linnaean Classification. Linnaeus, in his Philosophia
Botanica, divided systematists into heterodox and orthodox :
the former, those who classify plants by their roots, herbage, time
of flowering, place of growth, medical and economical uses, and
the like ; the latter, by the organs of fructification. It is remark-
able that all the orthodox or scientific classifications anterior to
Linnaeus made a primary division of the vegetable kingdom into
Trees and Herbs, referring the larger shrubs to the former and
the under-shrubs to the latter, an arrangement which began
with Theophrastus and was continued by Ray and Tournefort.
667. The three most important names in botanical taxonomy
anterior to Linnaeus are those of Cesalpini, Ray, and Tournefort.
Scientific botany commenced with the former, in Italy, in the
latter half of the sixteenth century. He first used the embryo and
its cotyledons in classification, distinguished differences in the in-
sertion of floral parts, and, indeed (excepting the primary division
into trees and herbs) , founded all principal characters upon the
organs of fructification, especially upon the fruit and seed.
Conrad Gesner of Zurich had somewhat earlier recognized this
principle, but Cesalpini first applied it.
668. A century later (1690-99) this principle was carried into
practice by Rivinus (a name latinized from Bachmann), of
BOTANICAL CLASSIFICATION. 333
Leipsic, in a wholly artificial classification founded on the corolla.
His contemporary in England, Robert Morison, somewhat earlier
began the publication of his great work, the Universal History
of Plants. In this was first attempted a grouping of plants into
what are now called natural orders ; and these were defined, some-
what loosely, some by their fruit, inflorescence, and flowers, others
by their stems, the nature of their juice, &c. But the two great
systcmatists of the time, who together laid the foundations of
modern scientific botaii}', were John Ray in England and Joseph
Pitton de Tournefort in France.
GG9. Ray's method of classification was sketched in 1682, and
was anterior to Tournefort's, but was amended and completed in
1703. The leading fault of both was the primary division into
trees and herbs. The great merit of Ray was his division of
herbs into Flowerless and Flowering, and the latter into Dicot}'le-
donous and Monocotyledonous. These great classes he divided
and subdivided, by characters taken from the organs of fructi-
fication, into what we should call natural orders or families, but
which he unfortunately called genera. He noted the coincidence
of nerved leaves with the monocotyledonous embryo, although
he did not notice that his first division of arborescent plants was
monocotyledonous ; and he had a clear apprehension of genera.
670. Tournefort's method was published in French in the
3'ear 1694, in Latin in 1700. It is more definite but more arti-
ficial than that of Ray, being founded like that of Rivinus almost
wholly upon modifications of the corolla, and it overlooked the dis-
tinction between monocotyledonous and dicotyledonous embryos.
Its great merit is that here genera, as we now understand them,
are first established and defined, and all the species then known
referred to them ; so that Tournefort was justly said b}' Linnaeus
to be the founder of genera. Ray ma}' be said to have indicated
the primary classes, Jussieu (in the next century) to have estab-
lished natural orders, and Tournefort to have given to botany
the first Genera Plan tarn m.
671. Linnaean Classification. Linnaeus, the great reformer of
botany in the eighteenth century, thoroughly re vised the principles
of classification, established genera and species upon a more scien-
tific basis, and, in designating species by a word instead of a
descriptive phrase, introduced binomial nomenclature. (704.)
He likewise established for the stamens, and indeed for the
pistils also, their supreme importance in classification (probably
without knowledge of the clear suggestion to this effect made
by Burckhard in a letter to Leibnitz, printed in 1702) ; their
functions, so long overlooked, being now ascertained. He also
334 TAXONOMY.
drew a clear and practical distinction between natural and arti-
ficial classifications (603) , and deferring all endeavors to make
the former available, except for genera, he devised a practical
substitute for it, as a key to the genera, viz. his celebrated
672. Sexual System, or arrangement of the genera under arti-
ficial classes and orders, founded upon the stamens and pistils.
Although now out of use, this artificial classification has been so
popular and influential, and has left so deep an impression upon
the science and especially upon the language of botany, that it
needs to be presented. The primary divisions are the classes,
twenty-four in number. But the 24th class, Cryptogamia, con-
sists of plants which have not stamens and pistils and conse-
quently no proper flowers, and is therefore the counterpart of the
remaining twenty-three classes, to which the corresponding name
of Phanerogamia or, in shorter form, Phamogamia (Phaenogamous
plants) has since been applied. These twent3'-three classes are
characterized by certain modifications and associations of the
stamens, and have substantive names, of Greek derivation, ex-
pressive of their character. The first eleven comprise all plants
with perfect (i. e. hermaphrodite) flowers, and with a definite
number of equal and unconnected stamens. They are distin-
guished by the absolute number of these organs, and are desig-
nated by names compounded of Greek numerals and the word
andria (from dvt'jo) , which is used metaphorically for stamen, as
follows :
Class 1. MONANDRIA includes all such plants with one stamen to the flower;
as in Hippuris.
2. DIANDRIA, those with two stamens, as in the Lilac.
3. TRIAXDRIA, with three stamens, as in the Valerian and Iris.
4. TETRANDRIA, with four stamens, as in the Scabious.
5. PENTANDRIA, with five stamens, the most frequent case.
6. HEXANDRIA, with six stamens, as in the Lily Family, &c.
7. HEPTANDRIA, with seven stamens, as in Horsechestnut.
8. OCTANDRI A, with eight stamens, as in Evening Primrose and Fuchsia.
9. ENNEANDRIA, with nine stamens, as in the Rhubarb.
10. DECANDRIA, with ten stamens, as in Rhododendron and Kalmia.
11. DODECANDRIA, with twelve stamens, as in Asarum and the Migno-
nette ; extended also to include those with from thirteen to nine-
teen stamens.
673. The two succeeding classes include plants with perfect
flowers having twenty or more unconnected stamens, which, in
12. ICOSANDRIA, are inserted on the calyx (perigynous), as in the Rose
Family; and in
13. POLYANDRIA, on the receptacle (hypogynous), as in the Buttercup,
Anemone, &c.
BOTANICAL CLASSIFICATION. 835
674. Their essential characters are not indicated by their
names : the former merely denoting that the stamens are twenty
in number; the latter, that they are numerous. The two fol-
lowing classes depend upon the relative length of the stamens,
namely :
14. DIDYNAMIA, including those with two long and two short stamens,
as in the majority of flowers with bilabiate corolla.
15. TETRADYXAMIA, those with four long and two short stamens, as in
flowers with cruciferous corolla.
675. These names signify in the former that two stamens, and
in the latter that four stamens, are most powerful. The four
succeeding are founded on the connection of the stamens, viz. :
16. MONADELPHIA (meaning a single fraternity), with the filaments
united in a single set, tube, or column, as in the Mallow.
17. DIADELPHIA (two fraternities), with the filaments united in two
sets or parcels, as in Corydalis and in many Leguminosse.
18. POLYADELPHIA (many fraternities), with the filaments united in
more than two sets or parcels, as in Hypericum.
19. SYNGENESIA (from Greek words signifying to grow together),
with the anthers united in a ring or tube, as in the Sunflower
and all Composite.
676. The next class, as its name denotes, is founded on the
union of the stamens to the style :
20. GYNANDRIA, with the stamens and styles consolidated, as in Cypri-
pedium and all the Orchis Family.
677. In the three following classes, the stamens and pistils
occupy separate blossoms :
21. MONCECIA (one household) includes all plants where the stamens
and pistils are in separate flowers on the same individual ; as in
the Oak and Chestnut.
22. DICECIA (two households), where they occupy separate flowers on
different individuals ; as in the Willow, Poplar, Moonseed, &c.
23. POLYGAMIA, where the stamens and pistils are separate in some
flowers and associated in others, either on the same or two or
three different plants ; as in most Maples.
678. The remaining class is essentially flowerless ; or rather its
organs of reproduction are more or less analogous to, but not
homologous with, stamens and pistils. But, although Linnaeus
suspected a sexuality in Ferns, Mosses, Algae, &c., there was no
proof of it in his day. So he named the class, containing these,
24. CRYPTOGAMIA, meaning clandestine marriage, the sexes, if existent,
hidden from view.
679. The characters of the classes may be presented at one
view, as in the subjoined table :
336
TAXONOMY.
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BOTANICAL CLASSIFICATION. 337
680. The orders, in the first thirteen classes of the Linnaean
artificial system, depend on the number of styles, or of the
stigmas when the styles are wanting ; and are named by Greek
numerals prefixed to the word gynia, used metaphorically for
pistil, as follows :
Order 1. MONOGYNIA, those with one style or sessile stigma to the flower.
2. DIGYNIA, those with two styles or sessile stigmas.
3. TRIGYNIA, those with three styles.
4. TETRAGYNIA, those with four styles.
6. PENTAGYNIA, those with five styles.
6. HEXAGYNIA, those with six styles.
7. HEPTAGYNIA, those with seven styles.
8. OCTOGYNIA, those with eight styles.
9. ENNEAGYNIA, those with nine styles.
10. DECAGYXIA, those with ten styles.
11. DODECAGYNIA, those with eleven or twelve styles.
12. POLYGYNIA, those with more than twelve styles.
681. The orders of class 14, Didynamia, are only two and are
founded on the pericarp, namely :
1. GYMNOSPERMIA, meaning seeds naked, the achenia-like fruits of a
4-parted pericarp having been taken for naked seeds.
2. ANGIOSPERMIA, with the seeds evidently in a seed-vessel or peri-
carp, /. e. the pericarp undivided.
682. The loth class, Tetrad}'namia, is also divided into
two orders, which are distinguished merely by the form of the
pod :
1. SILICULOSA; the fruit a silicle (561), or short pod.
2. SILIQUOSA ; fruit a silique (561), or more or less elongated pod.
683. The orders of the 16th, 17th, 18th, 20th, 21st, and 22d
classes depend merely on the number of stamens ; that is, on the
characters of the first thirteen classes, whose names they likewise
bear : as MONANDRIA, with one stamen, DIANDRIA, with two
stamens ; and so on.
684. The orders of the 19th class, Syngenesia, are six, namely :
1. POLYGAMIA /EQUALIS, where the flowers are in heads (the so-called
compound flower), and all hermaphrodite.
2. POLYGAMIA SUPERFLUA, the same as the last, except that the rays,
or marginal flowers of the head, are pistillate only.
3. POLYGAMIA FRUSTRANEA, those with the marginal flowers neutral,
the others perfect.
4 POLYGAMIA NECESSARIA, where the marginal flowers are pistillate
and fertile, and the central staminate and sterile.
5. POLYGAMIA SEGREGATA, where each flower of the head [or glom-
erule] has its own proper involucre.
6. MONOGAMIA, where solitary flowers, (that is, not united into a head)
have united anthers, as in Lobelia.
338 TAXONOMY.
685. The 23d class, Polygamia, has three orders, two of them
founded on the characters of the two preceding classes and
bearing their names, and the third named upon the same prin-
ciple, namely :
1. MONOSOIA, where both separated and perfect flowers are found in
the same plant.
2. DICECIA, where they occupy two different plants.
3. TRICECIA, where one individual bears the perfect, another the stami-
nate, and a third the pistillate flowers.
686. The orders of the 24th class, Cryptogamia, the Flower-
less Plants, are so man} 7 natural orders, and are not definable
by a single character. They are :
1. FILICES, the Ferns.
2. Musci, the Mosses.
3 ALGJE, which, as left by Linnaeus, comprised the Hepaticaa, Lichens,
&c., as well as the seaweeds.
4. FUNGI, Mushrooms, &c.
687. In its day, this artificial system well fulfilled its purpose,
and was preferred to all others on the score of facility and clefi-
niteness. Now no botanist would think of employing it, nor
would it be chosen for a key to genera, which was its only legiti-
mate use.
688. The Natural System was rightly appreciated by Linnaeus,
who pronounced it to be the first and last desideratum in syste-
matic botany ; and he early attempted to collocate most known
genera under natural orders (e. g. Piperitce, Palmce, Scitnmince,
Orchidece, Amentacece, &c., sixt}*-seven in number, including his
four cryptogamic orders) , but without definition or arrangement.
In his later years, he was unable to accomplish any thing more.
The difficult problem was taken up by Linnseus's contemporaiy
and correspondent, Bernard de Jussieu, who planted the botanic
garden at Trianon with plants grouped into natural orders, but
published nothing. His pupil, Adanson, who when a young
man lived for several years in Senegal, and who was as remark-
able for eccentricity as for erudition and ability, published in
1763, in his Families des Plantes, the first complete system of
natural orders. But he seems to have taken little from his
teacher, and with all his genius to have contributed little to the
advancement of the natural system.
689. Antoine Laurent de Jussieu, nephew of Bernard, has
been called the founder of the natural system of botany, and to
him more than to any other one person this honor may be
ascribed. In his Genera Plantamm secundum Ordines Natti-
rales disposita, 1789, natural orders of plants, one hundred
BOTANICAL CLASSIFICATION.
339
in number, were first established and defined by proper char-
acters, and nearly all known genera arranged under them. His
primary division of the Vegetable kingdom was into Acotyledones,
Monocotyledones, and Dicotyledones, adopted from Ray, with a
change which was no improvement. For his Acotyledones, the
Cryptogamia of Linnaeus, are the tk plants without flowers" of
Ray : they are, to be sure, destitute of cotyledons (though not in
the manner of Cuscuta) , because destitute of embryo altogether.
The Acotyledones forming his first class, Jussieu divided the
Monocotyledones into three classes upon single and artificial
characters, namely upon the insertion of the stamens, whether
hypogynous, perigynous, or epigynous ; and the Dicotyledones,
into eleven classes on similar characters, preceded by a division
into Apetalce, Monopetalce, Polypetalce, and Didinesirregulares, i. e.
first upon the character of the perianth, then upon the insertion
of the stamens or in Monopetalae of the corolla. The following
is the scheme :
Acotyledones t CLASS I.
Monocotyledones ....
Stamens hypogynous II.
perigynous III.
epigynous IV.
Dicotyledones
Apetalous
Monopetalous
Polypetalous
Diclinous (also Apetalous)
Stamens epigynous V.
perigynous VI.
hypogynous vn.
Corolla hypogynous VIII.
perigynous IX.
epigynous : anthers connate X.
epigynous : anthers separate XI.
Stamens epigynous XII.
hypogynous .... XIII.
perigynous XIV.
690. Auguste Pyrame DeCandolle was the next great syste-
matist. Reversing the order of Jussieu, who proceeded from the
lower or simpler to the higher or more complex forms, DeCan-
dolle began with the latter, the phsenogamous or flowering plants,
and with those having typically complete flowers. On account
of its convenience and the greater facilities for studying the higher
plants, this order has been commonly followed ever since. His
primary division on anatomical structure, into Vascular and
Cellular plants, was a backward step, confusing a portion of the
lower series with the higher ; and the duplicate names of Exo-
yence and Endogencf, appended to Dicotyledoneae and Monoco-
340 TAXONOMY.
tyledoneae, as it now appears should have been omitted. The
grades of the Candollean system superior to the orders, in their
final form, are mainty these :
])iv. I. VASCULAR (more properly PH^NOGAMOUS) PLANTS.
CLASS I. DICOTYLEDONOUS or EXOGENOUS.
Subclass I. THALAMIFLOROUS : petals (distinct) and stamens on the
torus, i. e. free.
II. CALYCIFLOROUS : petals (distinct or coalescent) and
stamens adnate to the calyx.
III. COROLLIFLOROUS : petals (mostly coalescent) not ad-
nate to calyx, bearing the stamens.
IV. MONOCHLAMYDEOUS : petals wanting.
CLASS II. MONOCOTYLEDONOUS or ENDOGENOUS. (No subclasses.)
Div. II. CELLULAR (more properly CRYPTOGAMOUS) PLANTS.
CLASS I. ^ETHEOGAMOUS : with sexual apparatus, and
Vascular tissue. (Eqnisetacece-Filices.)
Only cellular tissue. (Musci and Hepatlcce.)
CLASS II. AMPHIGAMOUS : destitute of sexual organs and of other than
cellular tissue. (Lickenes, Fungi, Algce.)
691. (>3 r ptogamous plants of all orders are now known to be
provided with sexes ; and the Jussiaean divisions of the Dico-
t3'ledones into Apetalce (including Diclines), Monopetalce, and
Polypetalce, are generally preferred to those of DeCandolle. Into
the present views of the classification of the Cryptogamia it is
unnecessary here to enter. Their general arrangement into
classes, &c., is not yet well settled, and the whole taxonomy
of the lower Cryptogams is at present in a state of transition.
602. John Lindley in successive attempts (between 1830 and
184o) variously modified, and in some few respects improved,
the Candollean arrangement. But, as neither his groupings of the
natural orders nor the new classes which he adopted have been
approved, his schemes need not be here presented. He must be
credited, however, with the first attempt to carry into effect a
suggestion made b}' Brown, that the orders should themselves
be disposed as far as possible into superior and strictly natural
groups. In Lincllej''s first attempt, such groups of two grades
were proposed, the lower called nixus (tendencies), the higher
cohorts. In his later and largest work, The Vegetable King-
dom, these were reduced to one, and the name of alliance was
coined. But this word has no good Latin equivalent, and the
term cohort (cohors) is preferred.
BOTANICAL CLASSIFICATION. 341
693. Robert Brown, next to Jussieu, did more than any other
botanist for the proper establishment and correct characterization
of natural orders. Having in the year 1827 published his dis-
coveiy of the g3'mnospermy of Coniferse and Cycadaceae, it
was in Lindley's works that this was first turned to proper
S3*stematic account by dividing the class of Dicotyledones into
two subclasses, the Angiospermce and the Gymnospermce. The
latter has been elevated by the vegetable paleontologists to the
rank of a class.
694. Stephen Ladislaus Endlicher, of Vienna, a contempo-
rary of Lindle}^ of less botanical genius, but of great erudition
and aptness for classification, brought out his complete Genera
Plantarum secundum Ordines Naturales disposita, between the
years 1836 and 1840. This elaborate work follows that of its
predecessor, Jussieu, in beginning with the lower series of plants
and ending with the higher. Its primary division is into two
regions: 1. Thallophyta, plants without proper axis of growth
(developing upward as stein and downward as root), no other
tissue than parenchyma, and (as was thought) no proper sexes.
This answers to the lower or Amphigamous Cellular plants of
DeCandolle. 2. Cormophyta, plants with an axis (stem and root),
with foliage, &c. The Cormoplryta, or plants of the higher
region, Endlicher divided into three great sections: 1. Aero-
brya, answering to the higher ^Etheogamous Cr} T ptogamia of
DeCandolle, with which was wrongly associated a group of root-
parasitic flowering plants (the Rhizanthese) which were fancied
to bear spores instead of embryo in their seed ; 2. Amphibrya,
which answer to Monocotyledones ; and, 3. Acramphibrya, which
answer to Dicotyledons. These last contain five cohorts: 1.
Gymnospermece ; 2. Apetalce ; 3. Gamopetalce (the Monopetalae
of Jussieu better named) ; 4. Dialypetalce (the Polypetalte of
Jussieu, &c.). The cohort in Endlicher's classification, it will
be seen, is a higher grade than that to which this name was
applied by Lindley in the more recent use. For the latter, i. e.
for the grade between these and the order, Eudlicher employed
the name of class.
695. Finally, the Genera Plantarum, now in course of pub-
lication by George Bentham and Joseph Dalton Hooker, adopts
in a general way the Candollean sequence of orders, with vari-
ous emendations ; divides the class of Dicotyledons into two sub-
classes, Angiospermous and Gymnospennous ; the former into
the Polypetalous, Gamopetalous, and Apetalous divisions ; and
the first of these into the Thalamiflorous, Disciflorous, and Caly-
ciflorous "series" (the middle one composed of the latter part
842 TAXONOMY.
of DeCandolle's Thalamiflorse with some of his Catyciflorse) ;
and under these the orders are arranged in cohorts, fifteen
cohorts in the Polypetalae, and ten under three " series" in th3
Gamopetalse. The remainder of this particular classification
has not yet appeared in print, although partly sketched b} T its
authors. It will generall}* be adopted in this country, with some
occasional minor modifications.
696. Various modifications have been from time to time pro-
posed. One of the best of them in principle is that initiated by
Adolphe Brongniart and adopted by many European botanists,
which, recognizing that most apetalous flowers are reductions or
degradations of polypetalous types, intercalates the Apetalae or
Monochlamydese among the Polypetalae. But this has never yet
been done in a satisfactory manner, or without sundering orders
which should stand in contiguit^y.
697. It should be borne in mind that the natural system of
botany is natural only in the constitution of its genera, tribes,
orders, &c., and in its grand divisions ; that its cohorts and the
like are as yet only tentative groupings ; and that the putting
together of any or all these parts in a system, and especially
in a lineal order, necessary as a lineal arrangement is, must
needs be largely artificial. So that even the best perfected
arrangements must always fail to give of themselves more than
an imperfect and considerably distorted reflection of the plan of
the vegetable kingdom, or even of our knowledge of it. 1
1 In the first place, the relationships of any group cannot always be
rightly estimated before all its members are known and their whole struct-
ure understood ; so that the views of botanists are liable to be modified with
the discoveries of every year. The discovery of a single plant, or of a point
of structure before misunderstood, has sometimes changed materially the
position of a considerable group in the system, and minor alterations are
continually made by our increasing knowledge. Then the groups which we
recognize, and distinguish as genera, tribes, orders, &c., are not always, and
perhaps not generally, completely circumscribed in nature, as we are obliged
to assume them to be in our classification. This might be expected from
the nature of the case. For the naturalist's groups, of whatever grade, are
not realities, but ideas. Their consideration involves questions, not of things,
between which absolute distinctions might be drawn, but of degrees of resem-
blance, which may be expected to present infinite gradations. Besides, al-
though the grades of affinity among species are most various, if not wholly
indefinite, the naturalist reduces them all to a few, and treats his genera,
tribes, &c., as equal units, or as distinguished by characters of about equal
value throughout, which is far from being the case. And in his works
he is obliged to arrange the groups he recognizes in a lineal series; but
each genus or order, &c., is very often about equally related to three or four
others : so that only a part of the relationship of plants can in any way be
indicated by a lineal arrangement.
BOTANICAL CLASSIFICATION. 343
698. Even the great classes cannot be arranged in a single
line, beginning with the highest Phaenogams and leaving the
lowest in contiguity with the higher Cryptogams. The Dicoty-
ledons take precedence of the Monocotyledons in rank. Yet a
part of them, the Gymnosperms, are much the lowest of all
known Phaenogams as regards simplicity of floral structure;
and through them only is a connection with the higher Cryp-
togams to be traced. The Monocotyledons stand upon an iso-
lated side line, and have no such simplified representatives.
In placing the latter class between the Dicotyledons and the
Acrogens, the chain of affinities is widely sundered. If, yield-
ing to a recent tendenc} r , we raise the G3~mnosperms to the
rank of a class, and place it between the Monocotyledons and
the Acrogens, then the much nearer relationship of Gymno-
sperms to Angiosperms through Gnetaceae and Loranthaceae is
not respected. (606, &c.)
699. Nor can the angiospermous Dicot}*ledons be disposed
lineally according to rank. The apetalous and achlanrydeous
must be the lowest. Some are evidently reduced forms of Poly-
pctalae or even of Gamopetalae : the greater part cannot without
violence be thrust into their ranks. The Gamopetalae, especially
those with much floral adnation, should represent the highest
type, the organs being at the same time complete and most dif-
ferentiated from the foliar state. If a natural series could be
formed, these would claim the highest place, with the Compositae
perhaps at their head. In the Candollean sequence, they occupy
the middle ; and the series begins, not without plausible reason,
with orders having generally complete blossoms, and such as
most freely and obviously manifest the homology of their organs
with leaves, then rises to those of greater and greater combi-
nation and complexity, and ends with those plants which, with
all their known relatives, are most degraded or simplified by
abortions and suppressions of parts which are represented in the
complete flower. These are low in structure, equally whether
we regard them as reduced forms of higher types, or as forms
which have never attained the full development and diversifica-
tion which distinguish the nobler orders.
700. Actual classifications, in their leading features and in
their extension to the cohorts, orders, &c., must be studied in
the systematic works where they are brought into use. In
these are the applications of the principles which are here
outlined. A separate volume of this text-book should illustrate
the structure, relations, and most important products of the
phaenogamous natural orders, as another is to illustrate the
344 TAXONOMY.
ciyptogamous orders. A S3~noptical view of the great divi-
sions only, as at present received and named, is appended.
Definitions and characters may be sought in the present and
preceding chapters.
SERIES I. PHLENOGAMOUS OR FLOWERING PLANTS.
CLASS! DICOTYLEDONS.
SUBCLASS I. ANGIOSPERMS.
Div. 1. POLYPETALOUS.
Div. 2. GAMOPETALOUS. (Monopetalous.)
Div 3. APETALOUS.
SUBCLASS II. GYMNOSPERMS.
CLASS II. MONOCOTYLEDONS.
Div. 1. SPADICEOUS.
Div. 2. PETALOIDEOUS.
Div. 3. GLUMACEOUS.
SERIES II. CRYPTOGAMOUS OR FLOWERLESS PLANTS.
CLASS III. ACROGENS.
Div 1. VASCULAR. (Ferns and their allies.)
Div. 2. CELLULAR. (Mosses and Liverworts.)
CLASS IV. THALLOGENS OR THALLOPHYTES.
PHYTOGRAPHY. 345
CHAPTER X.
PHYTOGRAPHY.
SECTION I. NOMENCLATURE.
701. PHYTOGRAPHY is the department of botany which relates to
the description of plants. This includes names and terms, also
figures and signs, as well as characters and detailed descriptions.
It comprises two sorts of names, one used to designate organs
or modifications of organs, the other to distinguish plants or
groups of plants. The former is Glossology or (to use the more
common but less proper word) Terminology. The latter is prop-
erly Nomenclature.
702. Names of Plants were at first only generic names. The
language of botany being Latin, and the plants which the old
herbalists knew being mostly European, their scientific names
were mainly adopted from the ancient Romans or, through Latin
literature, from the Greeks. Ex. Quercus, Prunus, Rosa, Rubus,
Trifolium ; and of Latinized Greek names, Agrostis, Aristolochia,
Colchicum, Melilotus. To the classical names others were
added from time to time ; as, from the Latin, Bidens, Convallaria,
Dentaria ; from the Greek, Anacardium, Glycyrrhiza, Loronthus,
&c. Some barbarous or outlandish names were early adopted,
such as Alhagi from the Arabs, and Adhatoda and Nelumbo from
India. These are mostly such as were or could be conformed to
Latin ; as Datura and Ribes from the Arabic, and later Thcea and
Coffcea. Of American aboriginal names, ffura, Guaiacum, and
Yucca are examples. Some ancient names of plants commem-
orated distinguished men. Ex. Asclepias, Euphorbia, Lysimachia,
Pceonia. Tournefort and his contemporaries resumed this
practice, and named plants in memoiy or in honor of distin-
guished botanists. Ex. Begonia, Bignonia, Ccesalpinia, Fuchsia,
Gerardin, Lobelia, Lonicera, Magnolia.
703. When among plants of the same name or kind different
species were known, these were distinguished by annexed epi-
thets. For example, among the Pines there were : Pinus syl-
vestris, vulgaris ; Pinus sylvestris, montana altera ; Pinus sylvestris,
montana tertia ; Pinus sylvestris maritima, conis fir miter ramis
adhcerentibus ; Pinus maritima minor ; Pinus maritima altera, &c.
346 PHYTOGRAPHY.
And as the number of known species increased, so did the
length of the phrases which were needed for their discrimina-
tion. These " differentiae," thus used as specific names (the
nomina specified of Linnaeus) , became extremely cumbrous. It
was about in the middle of his career that Linnaeus suggested
what he called trivial names (nomina triviulld) for the specific
name, consisting of a single word ; and in the Species Plan-
tarum, in 1753, he carried this idea into full effect in Botany.
The step was a simple one, but most important ; and Linnaeus
himself, who generally did not underrate his services to science,
seems hardly to have appreciated its practical value. 1
704. The Binomial Nomenclature in Natural History, thus
established, first separated the name of a plant or of an animal
from its diagnosis, descriptive phrase, or character, and reduced
the appellation to two words, the first that of its genus, the sec-
ond that of its species. The generic name very nearly answers to
the surname of a person, as Brown or Jones ; the specific answers
to the baptismal name, as John or James. Thus, Quercus alba is
the botanical appellation of the White Oak ; Quercus being that
of the genus, and alba, (white) that of a particular species ; while
the Red Oak is named Quercus rubra ; the Scarlet Oak, Quercus
coccinea; the Live Oak, Quercus virens ; the Bur Oak, Quercus
macrocarpa: Magnolia grandi flora is Large-flowered Magnolia;
M. macrophylla, Long-leaved Magnolia, and so on. The name of
the genus is a substantive, or at least is a word taken as a substan-
tive. That of a species is mostly an adjective adjunct, always
following the generic name and in the same gender. 2 This com-
bination of generic and specific name is the name of the plant. 3
705. By this system, not only is the name of the plant reduced
to two words, but a comparatively moderate number of words
serves for the complete designation of more than 120,000 plants, 4
1 Moreover, he may be said to have adopted rather than originated the
idea ; for single-worded specific names were used half a century previous
by Bachmann, alias Rivinus.
2 It is to be noted that the classical Latin names of trees are all feminine,
therefore Quercus alba, Pinus rigida, &c.
3 The name of a subgenus is sometimes written in between the two parts
of the plant's name, as Prunus (Padus) Viryiniuna. This is the name of the
plant and something more. In addition to the name of the species, that of
the variety or even subvariety is sometimes added.
4 Alphonse DeCandolle several years ago estimated the known species of
Flowering Plants at between 100,000 and 120,000. The larger number mny
perhaps include the higher orders of the Flowerless series. In the prese- t
state of our knowledge of the lower orders of Cryptogams, no close estimate
can be well formed of the actual number of species.
NOMENCLATURE. 347
in a manner which avoids confusion and need not overburden the
memory. The generic part of the name is peculiar to each
genus. The specific adjunct is not available for more than one
species of the same genus, but may be used in any other genus.
They are so widely thus employed that the number of specific
may not exceed that of generic appellations.
706. To render this system of nomenclature most serviceable
for the read}' identification of such numbers of plants or groups,
and for the clear and succinct presentation of or reference to
what is known and recorded of them, rules are indispensable,
and conformity to admitted rules is a manifest duty. Such rules
were s}'stematically formulated first by Linnaeus, in his Funda-
menta, Critica, and Philosophia Botanica, chiefly for generic
names, some of them being of the nature of laws, some rather of
recommendations. The most important of them remain in full
force, while many of the more particular rules restricting the
choice of names have been abandoned. The code was judi-
ciously revised (in his Theorie Elementaire) by DeCandolle
" who was ruled by the idea of having the law of priority prop-
erly respected," was critically considered by Lindley in his In-
troduction to Botany, and has of late been reformulated by
Alphonse DeCandolle under the sanction of a Botanical Congress
held at Paris in 18G7. 1
707. Rales for Naming Plants. These "should neither be
arbitrary nor imposed by authority. The}' must be founded on
considerations clear and forcible enough for every one to com-
prehend and be disposed to accept. The essential point in
nomenclature is to avoid or to reject the use of forms or names
1 Lois de la Nomenclature Botanique, etc., Geneva and Paris, 1867. In
the English edition, translated by Weddell : Laws of Botanical Nomenclature
adopted by the International Botanical Congress held at Paris in August,
1867, together with an Historical Introduction and a Commentary, London,
Reeve & Co., 1868. The Laws, simply, were reprinted in the American Journal
of Science and Arts, July, 1868. A few special points have been more recently
discussed by various critics, especially in the Bulletin of the Botanical Society
of France, and in that of the Royal Botanical Society of Belgium. See like-
wise American Journal of Science and Arts for September, 1870, and August,
1877 ; also, Bentham in Journal of the Linnean Society, xvii. 18P-108, in which
a just distinction is indicated between changing a well-established name and
giving a new name to a new plant. See American Journal of Science for
April, 1870.
Mention should also be made of Strickland's "Report of a Committee on
Nomenclature to the British Association in 1842, of Agassiz's classical preface
on the nomenclature of genera in his Nomenelator Zoologicus, and of Ball's
thorough and well-digested Report of the Committee on Zoological Nomen-
clature to the American Association for the Advancement of Science, 1877,
these dealing primarily with zoology.
348 PHYTOGKAPHY.
that may create error or ambiguity, or throw confusion into
science. Next in importance is the avoidance of any useless
introduction of new names. Other considerations, such as
absolute grammatical correctness, regularity or euphony of
names, a more or less prevailing custom, respect for persons,
&c., notwithstanding their undeniable importance, are relatively
accessory.' (Alph. DeCandolle, 1. c.)
708. The following are universal rules in scientific nomen-
clature :
1. Names must be in Latin or be Latinized. Those from the
Greek (which are more and more abundant, owing to the facility
of this language for compounding) take Latin form and termina-
tion. 1 Those from modern or other than classical languages
should at least have a Latin termination. 2 Hybrid names,
namely, those formed by the combination of two languages (at
least of Latin and Greek), should not be made. 3
2. For each plant or group there can be only one valid name,
and that always the most ancient, if it is tenable.
3. Consequently, no new name should be given to an old plant
or group, except for necessity. That a name ma}' be bettered
is no valid reason for changing it.
709. Names of Genera are substantive and singular, of one
word ; and the same name cannot be used for two genera of
plants. 4 They may be derived from any source whatever, from
1 Thus, words ending with the Greek os generally change it to us, and with
on to urn. A rule not always observed ; for while we have Epidendrum and
Oxydendrum, Linnaeus himself variously wrote Liriodendrum and Liriodendron,
Rlododendrum and Rhododendron ; and the Greek form now prevails.
2 In this as in other cases, some exceptions are well established by
custom, but they ought not to be extended. The rule as to Latinization is
restricted as respects orthography by the necessity of preserving modern
commemorative names in a recognizable form.
3 But we cannot change numerous old names for this fault, such as con-
volvuloides, fiimario>'des, ranunculoidcs, and scirpoides (though they ought to
have been convdmlind, ranunculina, and scirpina) ; and modern botanists have
not scrupled to append the expressive and convenient Greek term-aides (sig-
nifying likeness) to generic names not of classical origin. Ex. abutiloidcs,
bi.roides, darallioidcs, fuchsioides, f/enfia, Bursa Pdstoris, remain for sections and species, but not for gen-
era. When two words are confluent into one, they are not objectionable,
as Laiirorerasus, Carlemania (commemorating Charles. Li?man, Carolus Lpr
inanus), &c.
NOMENCLATURE. 349
prominent or peculiar character or appearance, from localities,
from the names of persons (especially of discoverers), from
indigenous or vulgar names, or even from arbitraiy combina-
tions of letters. Unmeaning names, if not in principle the best,
are never misleading. The main requisite is that they should
be euphonious, not too long, and that the}' should be adaptable
to the Latin tongue. Characteristic names, when possible, are
among the best ; such as Sanguinaria for an herb with red juice,
Htzmatoxylon for the Logwood tree, Lithospermum for a plant
with stony seeds (or seeming seeds) , Myosurus for a plant with
gyncecium resembling the tail of a mouse. Names of this sort
do not ftfw*ya hold out well ; for Chrysanthemum, so called from
its golden yellow blossoms, now has many white-flowered species,
Poly gala is wholly destitute of milk, and many species of Con-
volvulus do not twine. Neat anagrams are not bad, such as
Brown's Tellima for a genus nearly related to Mitella. Personal
generic names are wholly proper when dedicated to botanists,
especially to the discoverer of the plant, or to other naturalists,
or to persons who have furthered botanical investigation or
exploration. Ancient names of this kind have been mentioned,
also some of those which commemorate the earlier botanists.
(702.) At present, almost eveiy devotee of the science is thus
commemorated, from Linnaeus and Jussieu downward. In
forming such names, the name of the person, cleared of titles
and accessory particles (thus Candollea, not DecandoUea) , takes
the final -a or -ia and becomes feminine ; and its orthography is
preserved as far as possible, making only necessary concessions
to euphon}' and to the genius of the Latin language. 1
The Linnaean canon forbade the use of the same generic name in botany
and zoology, a rule now impossible to maintain. Perhaps we cannot pre-
vent the duplication of phsenogamous names in the lower Cryptogamia.
1 Thus, we may write Lescuria instead of Lesquereuxia, although Michauxia
is the form for the genus dedicated to Michaux, however pronounced. The
genus dedicated to Strangways is written Stranvcesia (although Strangwaysta
might have been tolerable) ; to Andrzeiowsky, Andreoskia ; to Leeuwenhoek,
Levenhookia (although the elder DeCandolle restored all the vowels), &c.
As specimens of overdone simplification, there is Gundelia, named for Gun-
delsheimer, and Goodenla, named for Bishop Goodenough, although Gundels-
heimera would not in these days be objected to, and Goodenovia is faultless.
Yet the names having been so introduced into the science should remain,
fixity being of more importance than perfection. Mistaken orthography
of the name itself may, however, be set right. Brown's Lecheruntltia is Les-
chenaultia, Nuttall's Wisteria (named after Dr. Wistar) is Wistaria. The
rule laid down in the code as drawn up by Alphonse DeCandolle is:
" When a name is drawn from a modern language, it is to be maintained
just as it was made, even in the case of the spelling having been misunder-
350 PHYTOGKAPHY.
710. The etymology of a new genus should always be given.
Of the Linnsean restrictions, one holds, viz. that the names of
genera are not to end in -oid>s, as man}* of the older names did.
71 1 . Names of Species are commonly and by preference adjec-
tives, agreeing with the name of the genus, and expressive of
some character, habit, mode or place of growth, time of flower-
ing, or commemorating the discoverer, first describer, or some
one otherwise connected with its history. Thus, in the genus
Ranunculus, R. bulbosus is named from the bulb-like crown or base
of the stem ; R. acris, from the acridity of the juice ; R. scele-
ratus (the accursed), in reference to the same property ; R. repens,
from the creeping habit of the stems ; R. pusillus, from general
insignificance ; R. aquatilis, from its growing in water ; R. ni-
vt^is, from living near eternal snow ; R. Pennsylvanicus, from
county or State whence it was first made known to botanists ;
R. Bonplandianus, in honor of Bonpland, one of the discoverers ;
and so on. More commonly, when a discoverer or investigator
of a species is commemorated in the name, this is a substantive,
in the genitive, as Ranunculus Nuttallu, i. e. the Ranunculus of
Nuttall, instead of R. Nuttallianns, the Nuttallian Ranunculus.
Yet the latter form is preferred when the species is named in
honor of some one who did not discover nor treat of it (which
should seldom be) ; but this distinction is a custom rather than
a rule, and the form of the commemorative name ma\ T be settled
by euphon}' or convenience. In any case, the personal name
should have a capital initial.
712. Many specific names are substantives, occasionally a
common substantive, as Stellnria nemorum (of the groves), Con-
volvulus sepium (of the hedges), Cassia pumilio (the dwarf) ;
more commonly it is a substantive proper name, and this usu-
ally an old generic name reduced to that of a species. Ex.
Ranunculus Flamm.ula, R. Thora, and R. Cymbalaria ; also //mo-
stood by the author, and justly deserving to be criticised." But this is
somewhat too absolute, since it is allowed that obvious errors in the con-
struction of names of Latin or Greek derivation may be corrected, provided
the change does not affect the initial letter or syllable, and that no ancient
names are to be disturbed.
The clause that forbids changes in the orthography of ancient names,
even to make them classical, is a very proper one. The botanical Latin
of Tournefort, Linnasus, Jussieu, arid their contemporaries, has by pre-
scription rights which botanists are bound to 'respect. Wherefore Pyrus
is the botanical name of the pear-tree, notwithstanding the classical Pirns.
So fan's, as a specific name for a smooth plant (and as distinguished from
lew's, a light or slight one), is fixed by long botanical use, although only
let-is is classical ; and it is unnecessary to change Ranunculus acris to R. acer.
NOMENCLATURE. 351
dendron Tulipifera, Rhus Toxicodendron, Dictamnus Fraxinella.
These proper specific names take a capital initial letter. 1 Rarely
such a name is in the genitive ; as Heterotheca Chrysopsidis, mean-
ing a species of Heterotheca with the aspect of a Chrysopsis.
713. Specific names should be of a single word. Some few
are compounded, as purpureo-cceruleum ; and some of ancient
origin (once quasi-generic) are of two words. Ex. Panicum
Crusgalli, Capsella Bursa pastor is, Taraxacum Dens leonis.
714. A specific name cannot stand alone. It is nothing
except as connected with the genus to which it pertains. A
Japonica by itself is wholly meaningless. A plant is named by
the mention of its generic appellation followed b}- the specific.
715. Names of Varieties. These are in all particulars like
specific names. Many are specific names reduced to a lowei-
rank. The varietal name is written after the specific, thus :
Ranunculus Flammala, var. reptans, and R. aquatiiis, var. tricho-
phyllus. Varieties of low grade need not be named. They ma3 r
be designated by numbers, or by the small letters of the Greek
alphabet, , J, &c. When the varieties are marked a and /3,
the first is supposed to be the type of the species, or both to
be equally included in the common character. But when the a
is not used, the varieties rank as deviations from the assumed
type of the species. Varieties of cultivation, half-breeds or
cross-breeds, and the like, should have only vernacular names,
at least not Latin ones such as may be confounded with true
botanical names.
716. Names of Hybrids are difficult to settle upon any com-
plete system. When of unknown or uncertain parentage, they
have been named in the manner of species, but distinguished by
the sign X prefixed. Ex. X Salix capreola. Hybrids of known
parentage are named by combining the names of the two pa-
rents, thus : S. purpureo X daphnoides, or X S. purpureo-daph-
noides, for a cross between S. purpurea and S. daphnoides, of
which the first supplied the pollen to fertilize the second. The
counterpart hybrid is X & daphnoideo-purpurea.
1 In respect to the initial of geographical specific names, being adjec-
tives, such as Americana, Canadensis, Virginiana, Europcea, Anglica, usage
governs, and this is divided. But the elder DeCandolle, who ruled in all
such matters in the preceding generation, always employed the capital in-
itial, and two generations of DeCandolle follow the example. Most English
authors until recently and some continental ones adopt this usage; and it
accords with the genius of the English language, in which we always write
European, British, American, &c., with a capital initial. Of late it is a usual
practice to write such geographical specific names with a small initial.
352 PHYTOGRAPHY.
717. The Fixation and Precision of Names. The name of a
plant is fixed by publication, and takes its date from the time
when it is thus made known to botanists.
718. A genus or other group is published when its name
and characters (or the differences between it and all other such
groups) are printed in some book, journal, or other adequate
vehicle of publication, which is placed on public sale, or in
some equivalent way is distributed among or within the reach
of botanists. A printed name without characters, and charac-
ters without name, do not amount to publication. 1
719. A species is not named unless it has assigned to it both
a generic and a specific name. It is not published until it is
made known, by name and characters (or by name along with
sufficient information as to its characteristics), in the manner
aforesaid. (718.) Adequate distribution, among botanists and
public herbaria, by sale or otherwise, of a collector's or distrib-
utor's specimens, accompanied by printed or autograph tickets,
bearing the date of the sale or distribution (that is, publication
by named Exsiccates in place of printed descriptions) , is held to
be tantamount to publication. 2
720. Characters, references to date and place of publication,
and the like, belong to bibliography or particular phytography,
not to nomenclature ; but proper identification of names requires
that the name of the author and the time and medium of pub-
lication should be taken into account. Anterior to the binomial
nomenclature, the botanical name of the common tall Buttercup
was "Ranunculus pratensis erectus acris" according to Bauhin,
in his Pinax, p. 179. Under the new nomenclature, which re-
duced the specific part of the plant's name to one word, this
became Ranunculus acris in Linnaeus, Species Plantarum (ed. 1),
p. 554 ; and a brief character gave its distinctions. In later
works it has been more fully described, in some illustrated by
figures. The citation of these works arranged in chronological
order (or in some order), with reference to volume, page, and
in some cases figures, is the bibliography of the plant. 3 A bot-
1 Names may be communicated, in manuscript or otherwise, by the pro-
pounder to an author who may make them known by publication ; but the
date of the genus or other group is that of actual publication.
2 This does not cover all the conditions of publication, since it does not
specify the characters (and the same may be said of a published figure,
with analyses) ; but, on the other hand, it conveys to the competent person
receiving the same all this information and more : so that it should carry
the rights of true publication as against any author to whom such names
are or should be known. That is, such arc not in the category of "unpub-
lished names," which generally ought to be left untouched.
8 For good examples of bibliography, see such detailed works as De-
NOMENCLATURE. 353
anist, in referring to this or any other plant, might cite any work
which describes it, or none at all. Ranunculus acris by itself,
as it happens, would lead to no ambiguity. Not so with many
names. For the accurate indication of the species, it is generally
needful, or highly convenient, to specify at least the name of
the author who first published the adopted appellation. So we
write Ranunculus acris, Linn., or L. , the abbreviated name of
Linnaeus. 1 Here we have the name of the plant, and the bibli-
ography reduced to its initial. To this, further citation and
other references may be added or not, as the particular case
requires. But, so far as citation or reference proceeds, it should
simply state the history correctly and clearly.
721. When a species is said to be of Linnaeus or DeCandolle
or Bentham, it is simply meant that the adopted name of the
plant (consisting of the generic and specific parts together) was
first published by this author. Some other author ma}' have
named it differently, and even earlier. The earlier name may
have been discarded because the specific portion of it was un-
tenable, either on account of preoccupation or for other valid
reasons. Or the later author may have differed from the earlier
in his views, and have referred the plant to some other genus.
As instances of the first, Euphorbia nemoralis, DarL, is a good
species, first named by Darlington in his Flora Cestrica. But
the name of Euphorbia nemoralis had already been applied to
and was the recognized name of a different species of the south
of Europe. Whereupon, as the North American species had
no other trivial name, a new one had to be given to it ; and it
was named E. DarUngtonii, in honor of the discoverer and first
describer. The common Milkweed of Atlantic North America
was named by Linnaeus Asclepias Syriaca. As this plant is not
indigenous to 'any part of the Old World, and does not at all
inhabit S}Tia, this trivial name is not merely faulty but false ;
so it was changed by Decaisnc into A. Cornuti, in commemora-
tion of an ante-Linnaean botanist who collected it in Canada and
gave the first account and figure of it. As an instance of the
second, take the pretty little vernal plant Anemone thalictroides,
L., meaning an Anemone resembling a Thalictrum. When it
was seen that the essential characters were rather those of Tha-
lictrum, the plant was placed in the latter genus. This was first
done in Michaux's Flora ; and so the accepted name is Tlialictrum
Candollc's Systcma Vegetabilium, and Sereno Watson's Bibliographical
Index to North American Botany, in the Smithsonian Miscellaneous Col-
lections.
1 For Abbreviations of Authors' Names, see 385.
354 PHYTOGRAPHY.
nnemonoides, Miclix., meaning an Anemone-like Thalictrum, and
Michaux is the authority for this name. The names which for
any reason are superseded become Synonyms. 1 (755.)
722. A later author may circumscribe a species or a genus
differently from the originator of the name. To a greater or
less extent, this must continuall}* happen in the course of time.
But u jRicinus cvmmums. Linn." stands unmoved by the sub-
sequent admission of various species (known or unknown to
Linnaeus) and the final reduction of all to one by a thorough
monographer. So does Silene Gallica, Linn., although S. quin-
quevulnera, Linn., of the same date, is reduced to it. There is
no sufficient reason for writing Myosolis, Brown, or Oynoglossum,
Brown, because this author restricted the limits of these genera ;
nor to write Gilia, Benth., because Bentham vastly extended
1 The synonymy is an essential part of the bibliography or scientific
history of a genus or species. But synonymous and admitted names ought
to be kept distinct. Keeping this principle in view, also the decisively
affirmed doctrine of the founder of our nomenclature, that the specific name
is a nullity apart from the generic (so that only the combination of the two
makes the name of the plant, as truly as the constituent halves make the
scissors), and bearing in mind the fundamental importance and absolute-
ness of the rule that no new names ought to be made where there are tena-
ble old ones, the student need not be misled by the confusing (however
specious) innovation countenanced by many zoologists and some botanists,
and which has of late years been very fully discussed.
The true rule is : " For the indication of the name or names of any group
to be accurate and complete, it is necessary to quote the author who first
published the name or combination of names in question." (A. DC ) Thus,
Lemtice thalictroides, Linn., fulfils the condition, except where a reference to
the work as well as the name of the originator of the name is demanded.
Then the citation would continue, " Spec. PL 312," and might be further ex-
tended. In the Flora of Michaux, this plant was treated as distinct from
Leontice in genus; and some botanists adopted this view, while others of
equal authority did not. Those who adopt Michaux 's genus name the plant
CaufopJi I/Hum ihnHctroides, Miclix.
Now some naturalists quote for the species the author who originated
the trivial appellation even when transferred to another genus. They
would adopt the genus Caulophyllum, yet write : Cnuhphyllum thalictroides,
Linn. Or else they would avoid direct falsification of the facts by adding
(sp.), this being explained to mean that the specific part of the name only
was given by Linnseus. Then, as this omits all mention of the original gen-
eric part of the name, others add this in a parenthesis, and write : " Canto-
pfit/ttnm thaJictroides (Linn, sub Leontice) Michx.," or " Caulophyllum (Michx.}
thalictroides, Linn, sub Leontice," or "Caulophyttum (Leontice, Linn.) thalictroides,
Micky." All such endeavprs to mix synonymy with nomenclature appear
to be faulty in principle and unwieldy in practice. In the most abbreviated
form, they state that which is not true : in the others, they impair the sim-
plicity and brevity of the binomial nomenclature. It is all but certain that,
if the genus Caulophyllum had been published in the lifetime of Linnaeus,
he would not have adopted it.
NOMENCLATURE. 355
the comprehension of this genus. Yet in their proper place
such changes may be indicated by " pro parte" or " char, muta-
tis" " excl. sp." and the like, useful qualifying statements,
but no part of the name.
723. Exactness requires that when a group is changed from
a higher to a lower rank, or the opposite, the name of tlia
author who made the change should be quoted. 1 He alone is
responsible for it. But this rule has only recently been strictly
observed.
724. In transferring a species from one genus to another, its
specific name must be preserved (with alteration of the gender,
if need be) , unless there is cogent reason to the contrary. It
must necessarily be changed when there is already in that genus
a species of the same name ; and then synonymous names of
the transferred species have their claim in order of date. But
whatever name is first emplo}*ed under the accepted genus, being
unobjectionable, should hold, even against an older unobjection-
able one coming from a w r rong genus. This is an application
of the stringent rule that no needless names should be created. 2
1 Thus, Potentilla Canadensis, L., var. simplex, Torr. $ Gray, and not of
Michaux, for it is the species P. simplex, Michx. Geum, subgen Sti/lipus,
Torr. fr Gray, not of Raf., for it is the genus Stylipus of Rafinesque, who
neither made the subgenus nor approved it. So, also, for the genus Labur-
num we write "Laburnum, Griseb ; " for even if it exactly corresponded with
Cytisus sect. Laburnum of DeCandolle, the latter is not a group of equiva-
lent rank.
But, as to genera and subgenera, this precision should not be insisted on
for times quite anterior to the recognition of such rules and of their need.
Speryularia began with Persoon as a subgenus in the year 1805, and this
date has been assigned to the genus, although it was taken up as such only
in 1819 by Presl and in 1824 by Bartling.
2 Thus, in the case of an older specific name being known, as that of
C/ii/o/tsis saligna, Don, recognized as B'ujnonia linear is, Car., though Don ought
to have adopted the latter trivial name, yet as he did not (and the rule was
not then really in force as now), there was no need for the introduction
of a third name, Chilopsis linearis, DC. " So, again, an Indian Grass was
first named and described by Willdenow as Coix arundinacea, then named
by Roxburgh as Coix barbata, and entered in Sprengel's Systema with
Willdenow's character as Coix Kceniyii. All these names were defective
as referring to a wrong genus. Brown corrected the error by creating the
new genus Chionachne, and selected Roxburgh's specific- name as the one
most generally known and the least liable to misinterpretation , and Brown's
Chionachne barbata is therefore the first correct name ; for which Thwaites
afterwards substituted Chionachne Kotnigii, an entirely new and useless name,
which falls by the law of priority. It should be well borne in mind that
every new name coined for an old plant, without affording any aid to
science, is only an additional impediment." Bentham (Notes on Euphorbi-
aceae, in Jour. Linn. Society, xvii. 197, 198, November, 1878). The following
356 PHYTOGRAPHY.
725. Names of Subgenera or of other sections of genera are
like those of genera ; indeed very man}' of them, and the most
fitting, are old generic names which have been comprehended
in the genus by reduction. Unlike genera and higher groups,
however, sections, when of Greek derivation, may properly take
the termination in -oides* and the typical section may bear the
name of the genus with the prefix Eu.' z Sections need not be
named at all, and only those of comparatively high rank should
is a farther extract from the same protest against the practice "of creating
a new name in order to combine an old specific with a new generic one : "
" In Ferns, the wanton multiplication of ill-defined or undefinable genera,
according to the varied fancies of special botanists, has had the effect of
placing the same species successively in several, sometimes seven or eight,
different genera ; and it is proposed to maintain for the specific appellation
the right of priority, not only in the genus alone in which it is placed, but
in the whole of the genera to which, rightly or wrongly, it has been referred.
This has been carried to such an extent as to give to the specific name a
general substantive aspect, as if the generic ones were mere adjuncts,
a serious encroachment on the beautiful simplicity of the Linnasan nomen-
clature ; and it is to be feared that there is a tendency in that direction in
phsenogamic botany. When a botanist dismembers an old genus, the rule
requires that he should strictly preserve the old specific names in his new
genera ; and, when he has wantonly and knowingly neglected this rule, it
may be right to correct him. But where a botanist has established what
he believes to be a new species, and has therefore given it a new name, the
changing of this name after it has got into general circulation, because it
has been discovered that some other botanist had previously published it
in a wrong genus, is only adding a synonym without any advantage what-
ever, and is not even restoring an old name ; for the specific adjective is
not of itself the name of a plant. ... A generic name is sufficiently in-
dicated by one substantive ; for no two genera in the vegetable kingdom
are allowed to have the same name ; but for a species the combination of
substantive and adjective is absolutely necessary, the two-worded specific
name is one and indivisible ; and combining the substantive of one with the
adjective of another is not preserving either of them, but creates an abso-
lutely new name, which ought not to stand unless the previous ones were
vicious in themselves, or preoccupied, or referred to a wrong genus. It is
probably from not perceiving the difference between making and changing
a name that the practice objected to has been adopted by some of the first
among recent botanists." Bentham, 1. c.
1 A genus could not properly have one of its sections called by its own
name with the addition of -aides or -opsis, as Asteroidcs or Astcropsis, for it
is senseless to declare that an Aster resembles an Aster ; but sectional names
of this composition may be excellent for sections of other genera, as ex-
pressing analogy or resemblance. Latin generic names used for sectional
ones properly take the addition of -ella, or -ina, or -astrum.
2 The prefix En (Greek for much, very, or true), prefixed to a generic
name of Greek origin, is the proper designation of the typical section of
that genus, meaning the group which should bear the generic name if such
genus were divided. The rule against hybrid names should in strictness
exclude this prefix from Latin names, but it has not always done so.
NOMENCLATURE. 357
have substantive names. Designations, however, are conven-
ient for lower sections ; and the name of a leading species
may be used, in the plural; as Aster, section Amelli, and sect.
Concinni. Subgenera need not agree in gender with the genus
the}' belong to. When written with the name of the plant,
the subgeneric name is parenthetically inserted between the
generic and specific appellation. Ex. Pyrus (Malus) coronaria.
72G. Names of Tribes, Orders, &c. The names of all groups
superior to genera are adjectives plural, and with few excep-
tions are the names of genera lengthened by some adjective
termination. Ex. From Rosa, Rosece, Rosacece, Rosales; from
Myrtus, Myrtea, Myrtacea, Myrtales ; from Berberis, Berber idece ;
from Tamarix, Tamaricinece ; from Salix, Salicece, Salicinece.
The substantive Plantce being understood, the groups are Rose-
ous, or Rosaceous, or Rosal plants, &c.
727. Tribal Names, and names of whatever grade between gen-
era and orders, are formed by adding to the root of a generic
name a final -ece. Ex. Rosece, Phaseolece, Antirrhinece, Oxalidece,
&c. Some sub tribes take the name of the tribe with the prefix
Eu, as Eiiphaseolece for that subtribe of the tribe Phaseolese
which comprises the representative genus Phaseolus. Tribal
names ma} T take the same prefix, as Euccesalpinece for the tribe
of the suborder Csesalpineae which contains the typical genus
Caesalpinia.
728. Ordinal Names are formed in the same way, but with a
preference for certain terminations which may denote their rank,
especially that of -acece, as Rosacece, Myrtacece, Cucurbitacece,
meaning Rosaceous, Myrtaceous, and Cucurbitaceous plants.
729. The names of what we now call natural orders, as
sketched or adopted by Linnaeus, were mostly descriptive, such
as Ensatce, Spathacece, Coronaria, Papilionacece, Coniferce, Amen-
tacece, Umbellate? ; but a few took their names from genera, as
Orchidece, Lillacece. Jussieu, with whom the system of natural
orders properly began, had no suborders, tribes, or any such gra-
dation of groups to deal with. His one hundred ordinal names
are some of them of the descriptive kind, as several of the above,
also Legumfnosce, Corymbiferce, &c. But the greater part are
simply plurals of generic names, such as Asparagi, Junci, Lilia,
Mxsce, Orchides, Lauri, Convohuli, Erica, Acera, Cacti. To a
few was given the lengthened termination in -ece, as Polygonea,
Solanece, Berberidece, Caryophyllece ; to some, the termination in
-acece, as in Cichoraccce, Campanulacece, Rubiacece, Ranuncu-
lacece, Malvacece, Tihacece, Gucurbitacece. Subsequent authors
have necessarily changed all names which were plurals of gen-
358 PHYTOGKAPHY.
era ; and the strongl}' prevalent tendency has been to give the
termination in -acecs to all such ordinal names, and to restrict
this termination to orders. Lindley insisted upon making this
an absolute rule even for names not formed from generic appel-
lations ; but this will not be adopted.
730. In the first place, several large orders which have been
known from the first by such characteristic names as Cruciferce,
Leguminosce (and its suborder Papilionacece) , Guttifera, Umbelii-
ferce, Compositce, Labiates, Cupuliftrce, and Coniferce, also Palmece
and GraminecB, Filices, and even Aroidece and Ficoidete, will retain
these appellations ; but no new ones of the kind will be made.
731. Also, names formed from genera which do not well take
the termination in -accce may be allowed as orders to retain their
natural form in -inece, -idece, -ariece, and the like. Ex. Tamaris-
cinece, Salicinece, Scrophularinece, Herberidece, Lentibulariece. We
ma}' prefer for the sake of uniformity to write Saiicacece, Berberi-
dacece, Lentibulariacece, and Scrophulariacece (as we should write
Violaceee), but this form cannot be insisted on. On the other
hand, a termination in -acece. has been allowed in the names of
certain tribes to avoid excessive iteration of vowels. Thus, for
the tribe of which Vernonia is the leading genus, authors write
Vernoniacece, to avoid Vernoniece, which ends with four vowels.
Spiraea and Staphylea are the types of tribes, for which the names,
if the} r followed the rule, would be Spirceece and Staphy/eece,
ending one in five the other in four consecutive vowels. Some
avoid this by writing Staphyleacece and Spirceacece. Others write
Staphylece, but this is only the plural of the generic name.
732. A few orders or other groups took their names long
ago from superseded generic names. Ex. Caryophyllacece or
Caryophyllece, Onagracece or Onagrariece, and Lentibulariece.
733. Names of Cohorts are distinguished by the termination
in -ales. This was proposed Iry Lindley, and is adopted by
Bentham and Hooker in the Genera Plantarum. Ex. Ranalcs,
Parietales, Malvales, Rosales* Passiflorales, &c., most of them
founded on the names of representative genera and orders.
Euphony requires some to take other terminations. Ex. Poly-
galince, CaryophylKncB.
734. Names of Classes and other great divisions are plurals,
either adjective or adjective nouns, expressive of the leading
character. Ex. Polypetalce, Gamopetalce, Apetalce ; Angiospermce
and Gymnospennce ; Dicotylcdones and Monocnfyhdones. The
names of the two great series or sub-kingdoms, following the
analogy of the Linnaean classes, end in -ia, and are Phcenogamia,
or Phanerogamia, and Cryptogamia.
GLOSSOLOGY. 359
SECTION II. GLOSSOLOGY OR TERMINOLOGY. 1
735. This is nomenclature as applied to organs or parts and
their modifications. The actual botanical terminology owes its
excellence in the first place to Linnaeus, and then to DeCandolle.
The Theorie Elementaire of A. P. DeCandolle (the first edition
of which was published in 1813) is still classical authority, and
until recently has received few additions as regards terms need
ful in phsenogamous botany.
736. The fundamental rule is that each organ or part shall
have a substantive name, and that modifications of organs shall
be designated by adjective terms. These names or terms should
be as precise as possible : each object ought to be known by only
one name, }*et sj^no^-ms are unavoidable ; and no term ought
to be used with two different meanings. The word flower, for
instance, must not be used for a cluster of flowers, however it
may imitate the appearance of one, nor for the corolla or other
portions of a flower. Still, some terms have to be used in two or
more senses, to be determined only by the connection, or else as
having both a special and a more general meaning. Leaf (fo-
lium) is a notable instance. A bract, to go no farther, is a sort
of leaf; and the imperfect stamens of a Catalpa-flower and
Pentstemon are stamens, although likewise called staminodia :
these are liable to be called sometimes by one, sometimes by the
other name. But, however frequent such ambiguities may be in
morphological treatment, they are usually avoidable in descriptive
botany, in which terms are held to their more special or partic-
ular sense. Yet no rule can absolutely determine whether leaf
or bract, bract or bractlet, is the proper term in many cases.
Moreover, substantive names must also be applied to certain
mere modifications of the same organ. In the same family, a
simple carpel, differently modified in fruiting, is an akene in a
Ranunculus, a follicle in Aquilegia, a berry in Hydrastis and
Actsea ; while in another family an additional line of dehiscence
makes it a legume. Moreover, in this latter family it is called a
legume when it is not dehiscent at all, and even when it becomes
a drupe ! Arbitrary rules cannot absolutely fix technical any
more than ordinaiy language.
737. Experience and judgment must determine what modifi-
cations of organs should be regarded as a kind, and bear sub-
1 Although the former is the better name, the latter is well established in
use as an English word, and perhaps it need not be objected to, inasmuch as
the Latin terminus comes from the Greek repp-a, of the same meaning.
860 PHYTOGRAPHY.
stantive instead of merely adjective names. But the former
should not be unnecessarily multiplied.
738. The classical language of scientific botany being Latin,
all the organs of plants and their principal diversities are desig-
nated by a Latin or Latinized name. Modern languages have
also their own names and terms. Greatly to its advantage,
English botanical terminology has adopted and incorporated
terms from the Latin and Greek, with slight changes, not obscur-
ing the identity, thus securing all their precision, and rendering
the simple botanical Latin of descriptions of easy acquisition to
the English student.
739. In a text-book like this, the principal names and terms
applied to organs and their leading modifications, as also those
which relate to their action (physiological terms) , or to our study
of them (didactic terms, such as phytography, phyllotax}', glos-
sology), are defined and illustrated in course. There remain
the more numerous and varied characteristic terms, chiefly adjec-
tives, applicable to more than one or to all organs, and which
compose the greater part of glossology. These, which DeCandolle
arranged systematically with much elaboration, may best be
reached by a glossary or dictionary, such as that at the end of this
volume, which comprises the substantive terms likewise.
740. From characteristic adjective terms are derived the
greater number of specific names of plants ; of which, therefore,
the glossary ma}' elucidate the meaning.
741. Capable as the existing system is, it cannot in single
words define all observed forms and grades, nor well avoid
various ambiguities of meaning. Some defects of the first kind
are remedied b}' combining with a Ir^phen two congruous terms
to denote an intermediate state. Ex. ovato-lanceolatus, or ovate-
lanceolate, for an outline between the two. Also a term may be
qualified by the prefix sub, in the sense of somewhat, as in sub-
rotundus, subcordatus (somewhat round or slight!}' heart-shaped),
or diminutives (such as integriusculus), or superlatives (integer-
rimits) or other strengthened forms (such as perangustus} may
be employed. Among terms of more than one form of meaning
are such as cah/cinus, which may mean, according to the context,
pertaining to the cal t yx, or of the appearance of calyx ; cymosus
may mean in c^mes, or bearing cymes, or in the manner of a
cyme ; and paleaceus may mean provided or beset with chaff, or
resembling chaff in texture. Often the form of the word should
distinguish the sense ; asfoliatits, furnished with leaves, foliosus,
with abundance of leaves, while foliareus may mean either bear-
ing leaves, or properly of leaf-like texture or appearance.
DESCRIPTION. 361
742. Absence of an organ or quality may be expressed by
means of a prefix with privative signification, as indehiscent, not
dehiscent, exaunulate, destitute of a ring, apetalous, without
petals. But the Greek privative should not be prefixed to
Latin words, nor the Latin sub to terms taken from the Greek.
743. When the Latin preposition ob is prefixed to an adjective
term, it means obversely ; thus obcordatus is cordate inversed.
that is, the broader end with its notch at the apex (instead of
the base) of the leaf or other plane organ.
SECTION III. DESCRIPTION.
744. Under this head may be conveniently comprised all that
relates to the form of the exposition, in botanical terms, of the
differences by which the species and groups of plants are distin-
guished and recorded, the structure exemplified, and the history
or bibliograplry indicated in systematic works or writings. Lin-
naeus, in the Philosophia Botanica, treated these topics under
the head of " Adumbrationes."
745. Descriptions may be full and general, comprising an
account of all that is known of the structure and conformation
of a plant or group, or rather all that is deemed worth recording,
or they may be restricted to what is thought most important. In
the former, the description is independent of all relative knowl-
edge, or takes no notice of relationship to other plants, or groups.
The latter intends to portray the species or group in its relations
to others, and to indicate the diiferences solely. Exhaustive
descriptions of the former kind are seldom drawn up, but partial
or supplementary ones are common. Descriptions of the latter
kind, when reduced to what is essential or differential, are termed
Characters, or the Character, of the group so described. There
are all gradations in practice between characters and descrip-
tions ; but the distinction should be maintained.
746. Characters are specific, generic, ordinal, &c. They are
the differentia, or marks which distinguish a group from any
related group of the same rank with which it ma)' properly be
compared. According to the occasion and purpose, they may
specify only the fewest particulars which will serve as a diag-
nosis, or the}' may be extended to all the known constant differ-
ences between two or more related species, genera, orders, &C. 1
1 The former would answer to what have been termed differential char-
acters, the latter to essential characters. Linnaeus divided (generic) characters
into factitious, essential, and natural; by the former denoting any difference
which may effectively distinguish between any two groups brought arti-
362 PHYTOGRAPHY,
What is now termed the specific character was the specific name
with Linnaeus and his predecessors ; what we call the specific,
Linnaeus called the trivial name. (703.)
747. Subordination of characters and the avoidance of vain
repetitions require that as far as possible regard being had to
the form of the work the ordinal character should contain
only what is needful to circumscribe it, and to exhibit clearly
its morpholog}* ; that the characters of tribes or other divisions
should not reassert any portion of the ordinal character, nor
the generic character that of the superior groups ; and so of the
sections and subdivisions of all grades down to the species.
Equally from the specific character should be excluded every
thing which belongs to the generic, or is common to its rela-
tives generally, or has been already specified in the section or
its subdivisions. So, likewise, of the varieties under the spe-
cies. This can be done only by so arranging the species as best
to exhibit their relationships, that is, by bringing together or
into proximity those of greatest resemblance in all respects,
or in the more important respects. What these are, and how a
just subordination of characters is to be apprehended, cannot
be taught by rules, but must be learned by experience and
from the critical study of the classical botanical works. No one
is competent to describe new plants without such study, and
without a clear conception of the position which a supposed
new specie^ should occup}' in its genus, or a genus in its order.
748. Characters of orders, genera, and of all intermediate
groups, are drawn almost without exception from the organs of
fructification. In the description, these parts are mostly taken
in order, beginning with the calyx and ending with the ovary,
the fruit, seed, embiyo. But, as to the orders, some writers pre-
fer to preface these proper characters with a general sketch of
those derived from the vegetation, which, albeit of less syste-
matic value generally, are often very characteristic of particular
families. Rubiacese, for example, are known by their opposite
entire and simple leaves and intervening stipules, along with a
few floral characters ; Sarraceniacese, b}- tubular or pitcher-like
leaves, along with a certain combination of a few other charac-
ficially together, as they might be in an artificial key, and as very unlike
genera often were in his sexual system ; by the second meaning the distinc-
tions, the fewer the better, which will separate a group from its nearest
relatives ; by the third, all real marks of difference, ?'. e. all afforded by the
organs of fructification, which only were taken into account for genera, &c.
Upon the construction of this natural character Linnaeus prided himself,
and justly. These are the characters in his Genera Plantarum.
CHARACTERS. 363
ters, and so on. Where brevity is aimed at, such external and
obvious characters, followed by a few diagnostic marks, ma}*
practically take the place of a full enumeration of particulars,
man}' of which may te common to other orders, though not in
the same combination. Generic characters alwa}*s commence
with the calyx or most external of the floral organs and proceed
to the ovary, thence to the fruit and seed, and end with subsi-
diary (but often no less diagnostic) particulars furnished by the
vegetation and mode of growth.
749. Detailed descriptions of species, as distinguished from
technical characters, commence with the root, and proceed in
order to the stem, leaves and their parts or appendages, inflor-
escence, bracts, flowers, calyx, corolla, stamens, with filament,
anther, and pollen, the disk, if any, g}*noeciiim and its parts,
ovules ; then the fruit, seed, albumen, if any, embryo and its
parts. But descriptions of this sort in most works and in ordi-
nary cases arc partial and subsidiary, comprising only certain
details supplementary to or in amplification of the character of
the species or genus. In condensed works, such description is
wholly omitted, or is reduced to a few specifications which do not
readily find their way into the character.
750. Specific characters usually follow the same order of
enumeration, from root to seed, so far as the several organs are
mentioned ; and in Latin the phrases are expressed in the abla-
tive case. But these particulars are often very conveniently
prefaced by statements applying to the whole plant rather than
to any one organ ; and these are given in the nominative, and
agree with the name in gender. 1
751. Linnaeus required that neither the essential character of
a genus, nor a specific character (his nomen specificum} , should
exceed twelve words. Latin characters take fewer words than
English. But this arbitrary rule is wholly out of date. Yet
such characters should be brief and diagnostic : otherwise, their
advantage is lost, and the distinction between them and descrip-
tions disappears. In monographs and floras, the desirable
brevity, or such as the case admits, is secured by proper group-
ing under a subordination of sections, subsections, and other
subdivisions. 2
1 Ex. " NEPETA CATARIA : erecta, elata, cano-pubescens ; foliis petio-
Jatis," etc. In English, these adjectives without any substantive expressed
will be seen to belong, as here, to " plant " or " herb " understood.
2 In the Synoptical Flora of North America, such a system of successive
divisions is thoroughly carried out. And, if the specific characters are by no
means short, it is mostly because nearly all separate descriptive matter is
364 PHYTOGRAPHY.
752. Punctuation. In proper descriptions, and in characton
of genera and of higher groups, the account of each organ forms
a separate sentence ; and in Latin the terms are in the nomina-
tive case, except subsidiary portions, which are often thrown
into the ablative. Excepting the latter part, the adjective terms
are separated by commas. A specific character is always in one
sentence. In Latin, its clauses are mainly in the ablative ; and
much diversity prevails as to the punctuation. 1 Subgeneric and
other sectional characters are commonly framed like those of
dispensed with: consequently various particulars are added to the char-
acter which do not strictly belong to it. In Bentham's great Flora Austra-
liensis, also in English, specific characters are replaced by a characteristic
synopsis at the head of each genus ; and a terse description under each
species completes the account. Moreover, Bcntham, in recent works, such
as his revision of the Genus Cassia, also that of the Mimoseae, which have
Latin characters, writes these in the nominative case and each member in a
separate sentence, in the descriptive form, abandoning the long-used abla-
tive form.
1 Linnaeus employed only the comma in the specific character, along with
a subsidiary use of the colon in a manner very unlike its ordinary use in
punctuation, making it a point of less value than the comma. Thus,
"CHENOPODIUM ALBUM foliis rhomboideo-triangularibus erosis postice
integris : summis oblongis, racemis erectis." Spec. PI. ed. 2, 310.
Here, while the two main members of the sentence are separated by a
comma, a subsidiary portion of the first member, relating to the uppermost
leaves, is separated by a colon. Linnaeus employed the colon in the same
way in generic characters. This anomalous usage is now abandoned. But
most authors have followed the Linnaean pattern in distinguishing the prin-
cipal members by commas only, so that these become the only points in the
specific character, however complicated that may be. Thus,
" RANUNCULUS ACRIS (Linn. Spec. 770) foliis pubescentibus subglabrisve
palmato-partitis, lobis inciso-dentatis acutis, summis linearibus, caule erecto
plurifloro subpubescente, pedunculis teretibus, calyce subvilloso, carpellis
nmcrone suberecto terminatis " DeCandolle, Prodromus, i. 36.
This is the punctuation throughout the Prodromus and in most contem-
porary systematic works. Its imperfection is shown in the above-cited speci-
men. The primary members of the sentence, which characterize the leaves,
stem, peduncles, calyx, and carpels, are distinguished by the same grade of
punctuation which serves for the parts of the first member, viz. the lobes
of the leaves, and for a still subordinate portion, viz. the form of the upper-
most lobes. This want of subordination is to be remedied by the use of
semicolons between the principal members, and of the commas only for the
secondary ones, a punctuation now not uncommon, and which is adopted
in the recent first volume of the Monograph iae Phanerogamarum of the De-
Candolles, which supplements the Prodromus. The portion of that volume
contributed by Dr. Masters better exemplifies this than does the rest of the
volume. For the latter sacrifices the advantage of the change by the inser-
tion of commas between each adjective of a continuous ablative phrase (as,
" /Smilax laitrifolia ; limbis foliorum oblongis vel ovato-oblongis, coriaceis,
3-5-nerviis, subtus pallidioribus," etc.), where they are generally deemed
PUNCTUATION, SYNONYMY. 365
genera. Or the members ma}' be united in one sentence, but in
that case the principal ones are best separated by colons.
753. Should a point intervene between the specific name and
that of the author cited? The practice varies. But, if the name
is Latin, the comma is superfluous ; for the abbreviated name of
the author is supposed to be in the genitive, and to read thus :
Ranunculus repens Linncei. Still, since when the author's name
is cited in full it is never written in the genitive, and since in
English the comma is normally required, it seems on the whob
proper to insert it.
754. In citations, the classical practice is to separate the refer-
ences from each other and from the name by periods ; thus,
"Anemone cylindrica, Gray, Ann. N. Y. Lye. 3. 221. Torr. &
Gray, Fl. 1. 113," 1 &c. It is becoming equally customary to
separate the several citations by semicolons, thus bringing all the
references under one name into one sentence. The bibliography
of a species or group of species which a describer or other author
has to refer to (with more or less fulness, according to the form
of his work) is to be sought partly under the admitted name,
and partly in the
755. Synonymy. This includes all other than the admitted
names. Ex. Hydropeltis of Michaux is a synonym of Brasenia of
Schreber, the latter being the earlier published name. Nectris
of Schreber is a synon^ym of Cabomba of Aublet, the latter hav-
ing prioritj*. Thalictrum anemonoides of Michaux has for syno-
n3'ms Anemone thalictroides of Linnaeus and of many subsequent
authors who followed him in referring this ambiguous plant to
Anemone (721) ; and also Syndesmon thalictroides of Hoffmannsegg
and Anemonella thalictroides of Spach, who proposed to consider
superfluous. The preferable punctuation of the character above-quoted
from the Prodromus would be
RANUNCULUS ACRIS (Linn.): foliis pubescentibus subglabrisve palmato-
partitis, lobis inciso-dentatis acutis, summis linearibus ; caule erecto pluri-
floro subpubescente ; pedunculis teretibus ; calyce subvilloso ; carpellis
mucrone suberecto terminatis.
The advantages of this style of punctuation will more and more appear,
when applied to less simple cases. Commas between the ablative adjectives
are superfluous and confusing.
In English characters, commas are required between the adjectives which
follow the noun. Rightly to express the subordination of characters, the plan
adopted in the Synoptical Flora of North America is recommended ; that is,
with colons separating the principal members, semicolons for subordinate
and dependent ones, and commas between the adjectives of the same noun.
1 See Watson's Bibliographical Index to North American Botany (where
this style is adopted) for a general model for the arrangement of synonymy
and citations.
366 PHYTOGliAPHY.
it an intermediate genus between Thalictmm and Anemone. In
systematic works, the specific character immediately follows the
name, and generally forms a part of the same sentence ; and is
followed first by citations of authors who have adopted the name,
and then by the synonymy, or as much of it as the plan of the
work calls for. The synonymous names and the references
under them should be cited in the order of their publication.
But, to economize space, all the authorities for the same name
are brought together into one sentence, and arranged according
to their date. Also, where the synonymy is not elaborately
displayed, the various synonyms of the same generic name are
usually placed in consecutive order/
756. Iconography. The leading and most essential citation is
that of the author by whom and the work in which a plant is
named and described, and also the work in which it is best char-
acterized. Among the characterizations, published figures hold
a prominent place. The citation of these is an important part
of the synonymy. The best botanical plates are those which
give detailed analyses of the parts of the flower, fruit, and seed,
displaying their structure.
757. Habitat and Station are recorded in a sentence or para-
graph following the name, character, and synonymy of a species.
The habitation is the place, district, or region at or within which
the plant is known to be indigenous, or to grow spontaneousl}'.
The complete habitat is the geographical range. The station is
the situation it affects, whether in water, in marshes, on shores,
on hills or mountains, in forests, on open plains, &c.
758. Discoverer, &c. To the habitat and station of newty
discovered, rare, or local plants should be appended the name
of the discoverer or the collectors by whom the species has be-
come known to science, at least when the plant is first published.
Date of discovery should also then be indicated.
759. Time of Blossoming should be recorded, either the month
or the season, to which ma}' be added that of the maturitj* of the
fruit. When the month or season is mentioned without farther
explanation, flowering-time is intended. In a flora, this may
sometimes be indicated under the genus for all the species.
In the flora of an extensive region, and in respect to species of
considerable range in latitude or longitude, the time of flowering
differs so widely at the extremes of the geographical range that
it cannot well be specified except in general terms, as spring,
summer, autumn, &c.
760. Etymology of Names. When a new generic name is pub-
lished, its origin and meaning should always be given, if the
ACCENTUATION, ABBREVIATIONS. 367
nature of the publication will allow it. So likewise of species,
except where the source or signification of the name is mani-
fest. This is commonly the case as respects most characteristic
specific names, and also those drawn from station, habitat, and
the like.
761. Accentuation of Names. The pronunciation of botanical
names is settled by the rules of Latin prosody. All that is
usually attempted in those botanical works which take this into
account is to mark the syllable upon which the principal accent
falls. This in words of two syllables is always the first ; in
words of three or more syllables, either the penult (the last sylla-
ble but one) or the antepenult (next preceding syllable) . When
the penult is a long syllable, it takes the accent ; when short, this
recedes to the antepenult. The accentuation may accordingly
be sufficiently indicated by marking the quantity of the penult,
either long as in Erica, or short as in Arbutus and Gladiolus. Or
else the accent may be marked by a proper sign, as Erica, Arbutus,
Gladiolus. An endeavor has been made to represent the longer
sound of the vowel b}' the grave accent-mark, as Erica, and the
short by the acute, as Gladiolus. But this plan is encumbered
with practical difficulties.
762. Abbreviations are required, both of the name of the au-
thor, when of more than one or two syllables, and of the titles
of the works cited. There are also the customar}' abbreviations
in the citation of volume, page, plate, &c., in which there is
nothing peculiar to botany.
763. The simple rule for the abbreviation of an author's name
is to abridge it of all but the first syllable and the first letter of the
following one (as Lam. for Lamarck, Hook, for Hooker), or the
first two letters following the vowel when both are consonants
(as Linn, for Linnaeus, Juss. for Jussieu, Rich, for Richard).
Sometimes more of the name must be given, in order to distin-
guish those beginning with the same syllable. So we write Michx.
to prevent confusion of the name Michaux with that of Micheli,
which, being the earlier, claims the abbreviation Mich., and
Bertol. to distinguish Bertoloni from Bertero. Sometimes a
much-used name of one syllable is abbreviated, as Br. (or R.
Br.} for Robert Brown. Initials or abbreviations of the bap-
tismal name are needed to distinguish botanists of the same
name ; as P. Browne in distinction from Robert Brown, Ach.
Rich., Adr. Jtiss., Alph. DC., to distinguish the younger from the
older Richard, Jussieu, and DeCandolle. Or, where father and
son, the abbreviation for the latter may be Juss.fL, Hook, f I.,
or Hook, f., &c. Certain, but very few, well-known and eminent
368 PHYTOGRAPHY.
names are abbreviated to a sign ; as L. for Linnaeus, DC. for
DeCandolle, HBK. for Humboldt, Bonpland, and Kunth, the
latter too long after ordinary abbreviation. 1 Care should be
taken to affix the period by which abbreviations may be dis-
tinguished from full names, such as Don, Ker, Blytt.
764. Abbreviations of titles of works follow the same rules as
those of names, or at least are in no wise peculiar in botany.
765. Abbreviations of the names of organs follow the same
rule : Cal. for calyx, Cor. for corolla, Stain, for stamen or
stamina, Pist. for pistillum or pistil, Fr. for fructus or fruit,
Per. for pericarpium or pericarp, Sem. for semen or seed, are the
most common. Hab. for habitat or geographical station, Herb.
for herbarium, Gen. for genus, Sp. or Spec, for species, Var. for
variety, and the like, every one will understand. But some
abbreviations which are common in botanical writings, at least
those in Latin, may need explanation to the elementary student.
A list of abbreviations is appended. See p. 390.
766. Signs. Under this head might be ranked such abbrevia-
tions as v. v. for vidi vivam, v. s. for vidi siccam, to note that the
writer has seen the plant, either alive or in a dried specimen ; or,
more particularly, v. s. s., when it is a spontaneous specimen
that has been examined in a dried state, and v. s. c., when it was
a cultivated specimen ; v. v. c., when the living plant was seen
in a garden only, and v. v. s., when the spontaneously growing
plant was seen alive. There are also proper signs, of which the
most common are those which indicate the sexes of blossoms,
the duration of a plant, and the like. Also the interrogation
point (?) used to express doubt ; the exclamation point (!) to
indicate the certainty that is given by the actual sight of an
authentic original specimen. See p. 391.
767. The marks used to indicate the subordination of sections
under a genus, or in the synoptical arrangement of genera, and
the like, are not settled by any fixed rule. An approved ar-
rangement is to employ the following marks in the given order,
* H -H- =. The first one, for sections of the highest order,
takes numerals after the sign. Ex. 1, and so on. When
1 As Alph. DeCandolle remarks, tlie proper abbreviation of the name lie
bears is Cand. But the form DC. was very early adopted by the first of
the illustrious name, and has be'.n continued for almost three quarters of a
century. Alphonse DeCandolle would prefer to write it D.C., but has not
adopted that mode, nor should we; for DC. and HBK. are convenient ab-
breviations reduced to signs. But such forms should not be increased. For
ordinary names they would be unintelligible.
Names which are not too long, and of which an abbreviation by the ordi-
nary rule is insufficient, such as Decaisne, should rather be written in full,
SIGNS, ETC. 369
such sections are followed by a substantive name, they are
equivalent to subgenera. 1 Ex. Phacelia, Juss., 1. Euphacelia,
i. e. the true or typical Phacelia ; 2. Cosmanthus, Gray, &c.
Sections next in rank to these are marked with asterisks, * for
the first, * * for the second, * * * for the third one of the same
rank. Divisions of these have the *- prefixed ; and so on in the
same way. Still farther subdivisions may be marked by the
small letters of the alphabet consecutively, a, b, c. When capital
letters are used for division marks, it is mostly for those of a
high grade.
7G8. Floras, Monographs, &c. A systematic work describing
in proper order the plants of a country or district is generally
called a Flora. A Flora of a small district takes the diminutive
name of a Florida. A universal work of the kind when it ex-
tends to the species is a System, Systema Vegetabilium or Sy sterna
lleyni Vegetabilis. The latest completed Systema Vegetabilium
is that of Sprengel (1825-1828), in five octavo volumes, on
a very condensed plan. A compendious Flora or S3'stema is
often termed a Prodromus, literally meaning a forerunner or
preliminary work. But, as even this is more than most bot-
anists are able to complete, the name of Prodromus is now
applied to works which are not intended to precede fuller ones
by the same author. The principal work of this kind is the Pro-
dromus Syst. Nat. Regni Vegetabilis, commenced by DeCandolle
in the j*ear 1824, continued b} r his son Alphonse DeCandolle
(aided by various botanists) to its close in 1873, down to vol.
xvii., or essentially twenty very compact octavo volumes, these
carrying the work only through the great class of Dicotyledones.
But the publication of the monocotyledonous orders has com-
menced in a series of Monographs (Monographic Phanerogam-
arum). A Monograph is a systematic account of all the species
of a genus, order, or other detached group.
769. Specimens of botanical characters and descriptions, cita-
tions, &c., illustrating this chapter, might be given here. But,
for those in Latin, the classical works of DeCandolle and others,
and for the genera those of Jussieu, Endlicher, Bentham and
Hooker, may be taken as models. In English, those of the
latter authors, and in the United States the better-known
writings of the present author, especially the later ones, may
be referred to.
1 DeCandolle in the Prodromus employed the word Sect. (Sectio) for
what answers to subgenus or at least to the highest grade of sections ; then
1, 2, &e., for the next grade below subgenus ; and then the asterisk, and
other marks.
24
370 PHYTOGRAPHY.
SECTION IV. SPECIMENS ; DIRECTIONS FOR THEIR EXAMINA-
TION, PRESERVATION, &c.
770. Implements. Those necessar3 T for the examination of
phaenogamous plants, Ferns, and the like, are a simple pocket
lens, a simple dissecting microscope ; also a sharp thin-bladed
knife and some needles of various fineness, mounted in han-
dles, for dissection.
771. Fora single hand lens, one magnifying only from four to
six diameters is the most useful. A doublet, or a parabolic lens
of Tolles, of about an inch focus, is better, but much more expen-
sive. The simple stage-microscope for dissection need have only
two lenses (doublets or otherwise) with large field arid good
definition, one of an inch and the other of about half inch focal
distance ; and a glass stage of at least an inch and a half in
diameter. A compound microscope is useful for all minute
investigation, and is essential in the study of vegetable anatomy
and of all lower ciyptogamic botan} T .
772. For making thin slices, a razor is the best knife ; for dis-
section on the stage of the simple microscope, beside needles,
small scalpels or some of the cutting instruments used by ocu-
lists are very convenient. But an expert hand is able to do
almost every thing with a common knife or scalpel and a pair
of mounted needles. Slender forceps are almost indispensable :
those made for the use of dentists are the best.
773. Analysis. In the examination of an unknown plant with
a view to its determination, its whole structure should be made
out, so far as the materials allow, before a step is taken to
ascertain its name and place in the sj'stem. In respect to the
stem, its duration and consistence and its internal structure,
whether exogenous or endogenous, are to be noted. As to the
foliage, the venation and the plvyllotaxy, also the presence or
absence of stipules, are most important. The anthotaxy or
inflorescence is to be examined and referred to its proper t}*pe.
In the flower, the numerical plan and symmetry, its ground-plan
and the nature of the deviations from the general or the family
type, are to be considered ; also the estivation or arrangement
of the parts in the bud, the character and extent of coalescence
and adnation ; the manner in which the anther is borne upon the
filament, and its place and mode of dehiscence, &c. Note also
whether, when the blossom is hermaphrodite, the anthers and the
stigmas mature at the same or at different periods. The placen-
tation and the character and position of the ovules should be
SPECIMENS. 371
determined. Two sections of the flower should be made : one of
them vertical and directly through the centre, in the manner of
Fig. 336-341, this will display the adnation, insertion, &c.,
of all the parts ; the other transverse and through the middle of
the ovaiy, also above the ovary when this is inferior, and if pos-
sible in the unopened but full-grown flower-bud ; this, among
other things, will bring to view the aestivation. (Fig. 351, 38G, 398,
&c.) Not rarely fruit and seeds are to be had at the same time,
or upon the same specimen, and these are equally to be investi-
gated. In fresh seeds, even those of minute size, the embryo
may almost always be extracted or brought to view under the
microscope, either by tearing away the seed-coat with needles or
by sections with a keen knife. When hard and dry, they have
only to be soaked or slightly boiled.
774. Diagrams and also sketches of the parts should be made,
such as those referred to in the foregoing paragraph. Such
diagrams can be drawn by any one with a little practice ; and
the}- ma}- be made to express the whole floral structure, even to
the coalescence and adnation. 1 But in the process of determina-
tion the student should beware of trusting wholly to his diagrams
and sketches without direct verification.
775. Dried specimens, when well prepared and in sufficient
abundance, in the hands of a skilled botanist are in most cases
but little inferior to fresh ones. When needed, flowers, or clus-
ters of blossoms, or fruits ma}' be detached and prepared for
examination and dissection by somewhat prolonged soaking in
warm water or by a short immersion in boiling water. This re-
stores flower-buds and small flowers and fruits, or their parts, to
a condition not essentially unlike the living state. Consequently,
the Herbarium or Hortus siccus of the botanist is to him more
essential than the botanical garden, important as that may be.
776. Herborizing. 2 The collector's outfit will essentially con-
sist of a Vasculum or botanical box, a Portfolio, a Trowel, a pocket
Lens, and a small but stoutly covered Note-book. Some use a
portfolio only, others the botanical box ; but on a long excursion
it is well to have both. The former is preferable in most cases,
except when specimens are collected for the immediate use of a
1 See Eichler's Bliithendiagramme (Leipsic, 1872, 1878), an admirable
work, which may serve as a model.
2 These articles, from paragraph 776 to 802 inclusive, were obligingly pre-
pared, at the author's request, by LYMAN H. HOYSRADT, of Pine Plains,
New York. They form an abstract or a new edition of a series of notes on
the subject which were published in the Bulletin of the Torrey Botanical
Club, in the year 1878.
372 PHYTOGKAPHY.
class. When well stocked with paper, it is of almost unlimited
capacity ; and most plants of delicate texture (as many of the
smaller aquatics, and those with fugacious or delicate corollas)
need to be consigned directly to the paper in which they are to
be pressed, and to be kept meanwhile under some pressure.
777. The Vasculum is very useful for holding plants that are
to be examined fresh, and for thick roots, large fruits, &c. It
is made of tin, and should be of oval-cylindrical shape, about 17
inches long and 4 by 6 inches wide, and provided with a light
strap to throw over the shoulder. The lid opens nearly the
whole length of one of the flat sides (15 by 4 inches, with one
fourth inch lap), is made to fit as close as possible, and fastens
by a simple spring catch. When no portfolio is used, a larger box
ma}* be required. Plants may be kept fresh in such a box for
many days. For a several-da3's excursion, when it is desirable to
bring home a large number of fresh plants, a tin chest, made some-
what after the pattern of an old-fashioned trunk, will be found
very convenient. It should be about 21 inches long, 10 inches
wide, and 10 inches high to the top of the convex and hinged
lid, which forms the whole top, and to which a handle is fitted.
778. A good form of Portfolio is made of two pieces of binder's
board covered with enamel cloth, and fastening together with two
long straps with buckles. Handles similar to those on a carpet-
bag may be attached for carrying. The usual size of portfolio
is 18 by 12 inches, but 16 by ll inches may be better, as
there would then be little danger of making specimens of too
great length for the herbarium. (784.) Or the back may be of
soft leather, an inch or so in width, and a light strap and buckle
at the front edge and at each end. The portfolio should contain
a good quantity of folded sheets of thin unsized paper, similar
to grocer's tea-paper, and of a size only a little smaller than the
sides of the portfolio. Very thin manilla paper, or what is so
called, is excellent for this purpose, being sufficiently bibulous
and rather strong.
779. The specimens as soon as gathered should be laid neatly
in these folded sheets (called specimen sheets) , and kept under a
moderate pressure in the portfolio. The sheets with the spec-
imens are afterwards transferred to the home press, but the
specimens should be left continuously in their sheets through all
the changing of driers, until cured. Indeed, the specimens may
well remain in the sheets after drying, until wanted for mounting
or for exchanging. For fine specimens, the use of this spectrum
paper is very important. Many plants are so extremely delicate
and sensitive that they will not bear the least Ijamlling without
HERBORIZATION. 373
curling and shrivelling, unless thus enclosed : also without these
sheets much time is lost in transferring small specimens one by
one from one drier to another in the drying process.
780. For digging up roots, bulbs, &c., a small and sharp
pointed triangular Trowel or stout knife will answer. One of
the best "diggers'* is made from a large file. Let a black-
smith bend the lower half of the blade to a gentle curve, so that
the point will be about an inch out of the true line. Grind off
the teeth and re-temper the blade. The total length with
handle, which is over one third, should be about twelve inches.
A leather case may be made for convenience of carriage. The
advantages of this strong tool are many. 1
781. A Note-book should be carried upon every excursion, in
which the station of rare plants, dates, colors, and various par-
ticulars which cannot be learned from the specimens, may
be recorded on the spot, instead of being left to uncertain
memory.
782. For most plants, the best time for collecting flowering
specimens is in the morning, soon after the dew has disappeared.
Vespertine flowers have to be secured earlier, or at nightfall.
783. Care should be taken to have the specimen of the proper
size, neither too small nor too large, and to comprise all that is
necessary for complete botanical illustration, flowers, fruit,
and leaves, both cauline and radical when possible. Inex-
perienced botanists suppose that a small sprig, containing a
flower or two with a few leaves, will answer all purposes as a
botanical specimen ; but later he comes to know better, and also
learns that the flower is only one of the component parts of a
specimen, and not always the most important one. In various
genera and orders, the fruit is the most distinguishing character-
istic, as with the Potamogetons, the Cruciferse, the Umbelliferae,
and the C3 r peraceae. With man}' plants the radical-leaves, with
others the character of the subterranean stem, whether a rootstock,
tuber, corm, or bulb, or of the root itself, whether annual, bien-
nial, or perennial, becomes important. Consequently, all the
organs have their value in an herbarium specimen, and each and
all should receive due consideration from the botanist when col-
lecting. Specimens may be often secured that exhibit both
1 [There is an English herborizing trowel of excellent quality, with blade
six or eight inches long, less than two inches wide, the sides slightly in-
curved, the stout shank an inch and a quarter wide, and one sixth of an
inch thick : this forms the whole back of the handle, the front of which is
a piece of lignum vitae riveted fast to the steel. It is nearly impossible to
break it.]
374 PHYTOGRAPHY.
flowers and fruit in the same plant, or fruit may be frequently
obtained from more advanced plants at the same time. Jf not,
fruit must be collected later, as in case of shrubs and trees, of
which generally only a branchlet with flowers, or with flowers
and leaves, can be gathered first. But subsequently the fruit
and mature leaves, should always be taken, if practicable, from
the same individual as the flowers. Of dioecious shrubs or
trees, like the Willows, each species should be represented by
four pieces : first, the sterile and fertile catkins will have to be
obtained, and the respective individuals marked, so that later
corresponding twigs with mature leaves, stipules, and fruit may
be gathered, and the specimens rightly matched.
784. A specimen should be so arranged as to be no larger
when pressed than can be neatly mounted on the herbarium
paper. A slender plant not over three feet in height should
generally be preserved entire, root and all. This can be done
by bending or partially breaking it at one, two, or three places,
and doubling so that the sections will not rest upon each other
in dr}*ing. If broken twice, it may be neatly arranged in the N
form when put in portfolio. Very large herbaceous plants will
have to be divided and the parts preserved separately, or, better,
take a suitable portion of the upper stem, having leaves, flowers,
and fruit, and a convenient part of the lower stem containing
radical leaves and with it sufficient root to show whether the
plant is an annual, biennial, or perennial. Thick stems, roots,
tubers, bulbs, and the like, should be divided or thinned down
with a knife, but in such a manner that the original shape can
be easily made out.
785. Carices should be always collected when the fruit is full-
grown, but not so ripe as to fall away. So also should other
Cyperaceae ; yet it is well to collect also earlier specimens of
these in flower. Grasses, on the other hand, should generally
be collected soon after they come into blossom. For when
mature the spikelets in many species break up and fall away in
drying. The culm, leaves, and root of Sedges and Grasses
should be preserved, as well as the inflorescence. The root is
no less important. Cespitose species should be so collected and
preserved as to show the tufted character. The culms of most
sedges and grasses act stubbornly when bent for arrangement
in portfolio or press, and are not disposed to stay in place. This
difficulty is promptly remedied by crushing with the teeth the
angles made by the bending. Or these may be thrust through
slits of paper. In diying Sedges and Grasses, very moderate
pressure should be employed.
HERBOR1ZATION. 375
786. Some aquatic plants (Algae especially) are so soft and
flaccid that, to secure them in their proper shape, they must be
placed in clear water and floated out by inserting beneath them
the paper on which they are to remain permanently, either the
regular mounting paper, or a thinner white paper which when
dry can be pasted on the herbarium sheet. If likely to adhere
to the sheet or drier above them in the press, a piece of oiled or
stearine paper may be laid directly on the specimens to prevent
their sticking. Also viscous or glutinous plants which are liable
to adhere to the sheets enclosing them may be sprinkled with
L/vcopodium spores, powdered soapstone, or some similar sub-
stance.
787. The name of the plant if known, but by all means the
locality and date of collection, with any other descriptive re-
marks regarded necessary, should be written on a ticket or on
the sheet when it is put into the press. Never omit to record
the time and place of collection, as a specimen of unknown date
and locality loses much of its value and interest.
788. Drying Specimens. The chief requisite for good herba-
rium specimens is the extraction of the moisture from the green
plant as rapidly as possible under a pressure which obviates
brittleness. This is to be affected by placing the thin sheets
containing the specimens between layers of bibulous paper, called
driers, and applying moderately strong pressure to the pile.
For driers nothing can be better than thick blotting paper,
except that it is too expensive, and the same may be said of
an English drying paper made for the purpose. Equally good
driers are made of the thick and felt-like brown paper which,
after saturation with coal-tar, is here largely used under the
clapboards of wooden houses and under slate-roofing. It is a
cheap material, and is to be obtained, cut into sheets of 18 by
12 inches. Or driers may be made of old newspapers or of any
soft wrapping paper, cut or folded to the proper size, and
stitched (very expeditiously by a sewing-machine), or joined by
eyelet paper-fasteners at two corners, in packages of a dozen or
more leaves to a drier. It is well to have a large supply of driers
and specimen-sheets ready for use.
789. A half dozen or more pieces of thin boards, 18 inches
long and 12 inches wide, should be provided. They are used
at the top and the bottom of the pile when pressing, and also
for dividing it into suitable sections, especially for separating
the packages of plants which were put into press at different
periods, and dividing up these packages themselves, if too large.
For the plants dry better in small sections and with the pressure
376 PHYTOGKAPHY.
evenly distributed. Hence it is best to have these sections not
over five or six inches in thickness, nor should the pile itself
be carried too high, never exceeding two feet. Painted binders'
boards may be used, instead of the common boards, to separate
interior divisions. Some botanists use a kind of lattice made of
two Ia3*ers of thin strips or laths, crossing each other. This is
said to allow free escape of the moisture by evaporation, and so
to accelerate drj'ing, as in the case of the wire press.
790. For 'giving pressure, various ways have been contrived.
The Screw-press is convenient and compact, but objectionable,
because it does not follow up the pressure as the plants shrink
in drying. This objection does not apply to the Lever-presses, but
they are usually unwiekty. Fortunately, one of the best forms
of the diying-press, as well as the simplest and cheapest, is
merely a board with weights placed on the top of the pile of
specimens. Here the pressure is continuous, constant!}' follow-
ing - the shrinkage of the plants. The weight on a pile should
vary from 25 to 100 pounds, according to the nature of the
specimens and the quantity in the press. On an average, 60
pounds is sufficient for most plants. If much greater pressure
is used, there is danger of crushing the more delicate parts of the
specimen, and thereb}' impairing its scientific value. For weights,
bars or masses of iron may be used, boxes filled with sand,
stones, and the like.
791. Specimens brought home in the botanical box must be
placed in such thin specimen-sheets as are used in portfolio.
In putting plants in specimen-sheets, whether in portfolio or
press, it is well to take some pains to spread out the specimens
neatly ; for a little care now may save much later labor. How-
ever, with most species, any carelessness in this respect can
be remedied at the first change of driers. But there are some
plants, previously referred to, so peculiarly sensitive that what-
ever adjustment the}' receive must be given at the time they are
first placed in their sheets.
792. Although plants can, if necessar} 7 , be kept fresh for several
da}'s in box or portfolio, on returning from a collecting trip they
should be transferred to the home press as early as possible. In
the transference, particular care should be taken to straighten
out and remove all folds and crumpling of the leaves, petals,
fronds, &c., and to arrange the specimen as naturally as possi-
ble, so as to show the proper habit. Both sides of the flowers
and leaves should be exhibited. Plants that were put directly
into press should receive this special attention at the first
change of driers, which on this account should be made within
HERBORIZAT1ON. 377
several hours afterwards. The stubbornness and elastic! ty, so
troublesome in specimens when first put in, will then have
mostly disappeared, and the whole specimen will be found suffi-
ciently flaccid to have every part stay as arranged. If this first
change is deferred longer than ten or twelve hours, the speci-
mens of man}' species become too dry for making the alterations
required. At this time small pieces of bibulous paper may be
placed between leaves, or other portions of the plant which over-
lap, to prevent moulding or discoloration, and to hasten drying.
It is well to change these fragments of paper with the driers for the
first day or two : afterwards they may remain with no detriment.
793. To have the specimens retain their natural color and
general appearance, the}* should be dried as rapidly as possible ;
and this result is best secured by frequent changing of the driers.
These should be changed at least once a day for the first four or
five days, and afterwards ever}* other da}', until the specimens
are thoroughly dried. But a marked improvement in the speci-
mens will result from more frequent changes during the first
day or two. The first day with Grasses, Sedges, and their allies,
and the first two days with most other plants, are of more impor-
tance than all the subsequent time. As an experienced collector
declares : " Two or three changes of the driers during the first
twenty-four hours will accomplish more than a dozen changes
after the lapse of several days. The most perfect preservation
of the beautiful colors of some Orchids has been effected by
heating the driers and changing them every two hours during
the first day."
794. Heated driers are very efficient ; and the best mode of
heating is to expose them to the sunshine, and bringing them in
hot to make the change at once, or as soon as possible.
795. The number of driers interposed between the specimen-
sheets should depend upon the plants and the frequency of the
changes : two will suffice when the driers are changed very often ;
but more must be employed when the plants are thick and succu-
lent. Uniform pressure ma}' be secured with large and coarse
plants by placing strips of pasteboard or pieces of cotton-bat-
ting about the sides of the package. Ringlets of cotton may be
placed about some of the larger flower-heads of the Composite, &c.
796. The time required to dry specimens varies with different
species and with the season : it depends also on the frequency
of the changes and the temperature of the driers. By changing
daily, the time is usually from four or five days to a week. But,
with two changes a day for the first day or two and with heated
driers, the process may be completed in half the usual time, and
378 PHYTOGEAPHY.
the specimens will be in much finer condition. An experienced
collector has no difficult}" in ascertaining whether a plant is com-
pletely cured or not, while to a novice it is often a matter of
uncertainty. A thoroughly dried plant can be usually told by
its peculiar hay-like rattle when disturbed ; also by placing the
plant against the cheek. If there is a sensation of coolness, the
plant is still moist.
797. If the thick leaves of fleshy plants are immersed for
a few moments in hot water, the period of desiccation will be
greatly hastened ; but they frequently turn dark as a conse-
quence of the immersion. The drying of such plants, and
particularly of the Monocotyledons, ma}' be advantageously ex-
pedited by placing them between several driers and ironing them
with hot irons. Small plants may be very neatly dried in old
books. Very beautiful specimens may be made by placing the
plant in a tall and narrow vessel, and pouring over it a sufficient
quantity of clean and dry sand. When the moisture is absorbed,
it may be flattened in a press.
798. In shifting the driers of a collection, place the package
to be changed at the left hand on the table or counter, the new
pile in front with its length parallel to the person, a position
the most favorable for giving any needed attention in arranging
specimens, while fresh driers maybe placed at the right hand,
or be}'ond the pile in front. Thus arranged, the sheets of speci-
mens can be rapidly shifted into their fresh driers.
799. The moist driers may be spread out in the sunshine to
dry, or strung on a line in a warm room, or in the open air, if
not too windy. Very moist driers may be thoroughly dried
within an hour, if spread in the hot sunshine. In inclement
weather, they must be dried by the fire.
800. To recapitulate the most important points in good speci-
men-making : Use specimen-sheets to hold the plants undis-
turbed during the whole process of drj'ing : use plent} T of the
most bibulous driers, sun-dried and heated when practicable : do
not make the piles too large : make the first shift of driers within
a few hours, at that time making all needed adjustment of the
flaccid specimens: change the driers twice a day for the first
day or two.
801. For collecting and preserving specimens on a journey,
or when moving from place to place, some modification of the
station ary press is requisite. The Travelling-press must be porta-
ble : accordingly the pressure is applied by strong leather straps
with buckles. There should be three straps, one girding the
package around each end, and one lengthwise. The top and
HERBORIZATION. 379
bottom, if of thin boards, must be cleated. or compounded of
wood with the grain in opposite directions, or ver} r stout binder's
board or trunk-board may be advantageously used. This should
be covered with coarse cotton or linen cloth, glued fast and well
painted. While stationary, the pressure may be given by means
of weights when more convenient.
802. The Wire press, now much in use, is a press of this porta-
ble kind, in which the boards are replaced by sheets of wire net-
ting, with wide meshes, and surrounded and strengthened by a
strong but light iron border. Straps with buckles are used to
hold the parts and contents together and to apply the pressure,
as in the ordinary travelling-press. Besides its portability, the
advantages of such wire presses are that, in a small way, they
may serve both as portfolio for collecting and as press for dry-
ing ; also that, as the drying takes place mainly by evaporation
instead of absorption, much less paper is required, and the trouble
of changing the driers is saved. 1 In fair weather, the press filled
with plants may be hung in the wind or sunshine, in foul weather
near a fire. The disadvantage is that specimens dried in this
way are apt to be brittle. To use this s3 T stem advantageously,
the botanist should have at least two such presses in operation,
one for collecting, while the other is in use for drying.
803. Poisoning. Dried specimens are liable to the depreda-
tions of certain insects, especially of their larvae. The principal
pest is a small brown beetle, Anobium paniceum, L. ; the perfect
insect does considerable damage, the larva vastly more. Plants
with milky juice, such as Asclepiadeae, Apocynaceae, and Eu-
phorbiaceae, those containing bitter principles, such as Gentians
and Willows, and generally such plants or such organs as con-
tain much protoplasm or azotized matter, are most subject to
attack. Ranunculaceae, Umbelliferae, and Compositae are seldom
spared ; while Labiatae mostly escape, probably on account of the
volatile oil which they contain. Even Ferns are liable to have the
parts of fructification eaten away. To a certain extent, the im-
pregnation of the herbarium-cases with camphor, naphthaline,
or strong-scented oils, may exclude the vermin. But safety is
secured only by poisoning.
804. The proper poison is corrosive sublimate, dissolved in
strong (95 per cent.) alcohol. Drop into the alcohol as much cor-
1 Prof. A. Wood seems to have been the first to call the attention of
American botanists to this system, which he has earnestly advocated.
An improved form of this wire press, well adapted both for collecting
and pressing in moderate quantity, is made and sold, at a small price, by
Paul Roessler, optician, at New Haven, Connecticut.
380 PHYTOGKAPHY.
rosive sublimate as it will take up, then add a trifle more of alcohol,
so as to keep the solution just below the point of saturation.
The stronger the solution the better, except that, at full satura-
tion and where copiously- used, an efflorescence may sometimes be
left on the surface of the poisoned specimens upon the evapora-
tion of the alcohol. Some add to the solution some carbolic
acid, at the rate of a fluid ounce to each quart of alcohol. The
solution may be applied with a soft brush (one with no metal in
its fastening) ,' or by a dropping bottle, or even the specimens
may be dipped in the solution placed in a flat porcelain dish.
The brush (using a pretty large and soft one) is the most con-
venient and efficient. The moistened specimens should be placed
between driers and in shallow piles until the alcohol evaporates.
805. Thoroughly poison all specimens before admitting them
to the herbarium. It is well to poison all specimens whatever,
as soon as they are made or at the close of the botanizing sea-
son, as well those intended for exchanges as for the collector's
own herbarium.
806. Keep all specimens between sheets of paper, or within
folded sheets, not too crowded or overlaid, away from dust, and
in a perfectly dry place, so as to avoid mould. When attacked
b}' mould, the corrosive-sublimate solution should be applied.
A properly dried specimen, dulj' cared for, should be as lasting
as the paper which holds it.
807. The Herbarium, called by the earlier botanists Hortus
Siccus, is a collection of dried specimens, named and S3'stemat-
ically arranged. It is indispensable to the working sj'stematic
botanist, and every devotee of botany should possess, or have
access to an herbarium containing representatives of the plants
of the immediate vicinity or district, if not of the whole country.
Or an herbarium may be restricted to a particular family of
plants, made the object of special study. A general herbarium
should contain specimens representing all the natural orders and
as many of their genera and species as possible.
808. The form of the herbarium as to the size of its sheets is
considerably variable. That of Linnaeus is of the size of foolscap
paper : this would now be universally regarded as much too
small. The principal British herbaria adopt the size of 16 J by
10 J inches, which is rather too narrow, rarely permitting two
specimens of the same species of any considerable size to be
placed side by side on the same sheet. In the United States,
16^ inches in length by llf in width is adopted ; that is, for the
genus-covers, the species-paper being a quarter of an inch nar-
rower.
THE HERBARIUM. 381
809. The specimens representing each species may either be
laid within a doubled sheet, loosely (as in some European her-
baria), or fastened in place by narrow slips of gummed paper
(which is much better), or else they maybe glued bodily to
single sheets of strong and stiff white paper.
810. The former is an excellent plan for a limited collection.
It is an advantage that a specimen can be taken up and examined
on all sides ; also, that indifferent specimens can at an} T time be
exchanged for better ones. But a large herbarium on this plan
becomes cumbrous and inconvenient for ready reference and
comparison.
811. The best plan in a large herbarium, and one much to be
consulted, is to attach the specimens completely, by an} 7 kind
of strong and light-colored glue, to single sheets, or rather half
sheets. The specimens are thus safe from injury under reason-
able handling, and can be turned over and examined with as
much facility as a series of maps or engravings. The species-
paper should be of writing-paper stock, or of equal firmness, of
compact texture, well sized and calendered, and of a weight in
size of 16 by 11 1 inches of about 18 pounds to the ream of
480 flat sheets. The paper should be furnished square-cut on
all sides, in the manner of "flat cap." Stiffness is the great
desideratum.
812. In no case should more than one species be knowingly
attached to the same sheet. But of very many species there will
be room for more than one specimen. And specimens from dif-
ferent localities, of different forms, and in various stages of
flowering and fructification, are always desirable. The full name
of the plant should be written at the lower right-hand corner of
the sheet, or a ticket should there be attached by glue or traga-
canth paste. Each specimen should have its ticket, similarly
attached, or a memorandum upon the sheet, indicating the hab-
itat or the special locality, date of collection, name of collector,
and any other desirable information which the specimens them-
selves do not furnish. When there are loose flowers or fruits, or
when any of these have been detached for dissection and micro-
scopical investigation, it is well to preserve them, placing them
in little paper pockets or envelopes and pasting these upon the
sheet close to the specimen to which they pertain. Sketches of
parts dissected may be drawn upon the sheet. Notes and mem-
oranda received with the specimen or too extended to be entered
upon the sheet may be folded, inserted in such envelopes, and
made fast to the sheet. Many botanical collections are distrib-
uted with printed tickets. These, and all authenticating tickets
682 PHYTOQKAPHY.
or notes, should be attached to the sheet near to the specimen
they belong to. In view of this, printed and written tickets
should be of small size. 1 A ticket which exceeds four by two
1 All printing on an herbarium ticket should be in plain type ; and fancy
borders, uselessly occupying room, should be avoided. If any border is
thought needful, it should be of plain lines. It is not desirable to parcel out
the space on a ticket with separate lines and headings for habitat, date of
collection, time of flowering or fruiting, name of collector, and the like.
These particulars may conveniently be entered at the bottom or top of the
ticket, as may be convenient, leaving the rest of the space free for the name
of the plant, the authority, and perhaps a synonym.
Tickets for specimens distributed among other botanists may well have a
head-line indicating the source, such as " EX COLL. c. c. PARRY," or " EX HERB."
or, in English, " FROM THE HERBARIUM OF " the botanist who communicates
the specimen. The following may serve as an example of a simple ticket
for the sending out of dried specimens, and of the way in which the ticket
may be filled out with the name of the plant, its habitat and station, name
of collector and time of collection.
EX HERB. A. GRAY.
J. G. Agardh, son.
Bess.
Besser.
Ait.
Aiton.
Bieb.
MarschallvonBieberstein.
AIL
Allioni.
Bigd.
Jacob Bigelow.
Amm.
Amman.
Bisch.
Bischoff.
Anders.
Andersson of Stockholm.
Bcehm.
Boehmer.
Andr.
Andrews.
Boerh.
Boerhaave.
Andrz.
Andrzejowski.
Boiss.
Boissier.
Aresch.
Areschoug.
Boland.
Bolander.
Am.
Arnott.
Bong.
Bongard.
Arrh.
Arrhenius.
Bonpl.
Bonpland.
Asch.
Ascherson.
Bork.
Borkhausen.
Aubl.
Aublet.
Borsz.
Borszcow.
Brack.
Wm. D. Brackenridge.
Bab.
Babington.
Brebis.
Brebisson.
Bail.
Baillon.
Bref.
Brefeld.
Balb.
Balbis.
Brew, fr \
W. H. Brewer & Sereno
Baldw.
Baldwin.
Wats, J
Watson.
Balf.
Balfour.
Brid.
Bridel.
Barn.
Barneoud.
Brong.
Brongniart.
Barr.
Barrelier.
Brot.
Brotero.
Bart.
Benj. Smith Barton.
Brouss.
Broussonet.
W Bart.
W m - P. C. Barton, nephew.
Br., R. Br.
Kobert Brown.
Bartr.
John Bartram.
P.Br.
Patrick Browne.
Bartr.f.
Wm. Bartram.
Brunf.
Brunfels.
Bauh.
Bauhin.
Buckl
Buckley.
Beauv.
Palisot de Beauvois.
Bull.
Bulliard.
Benj.
Benjamin.
Burm.
Burman.
Benn.
J J. Bennett.
Buxb.
Buxbaum.
A. Benn.
A. W. Bennett.
Benth.
Bentham.
Cam.
Camerarius.
Berg.
Berk.
Bergius.
M. J. Berkeley.
Camb. )
Cambes. }
Cambessedes.
386
ABBREVIATIONS.
Campd. -
= Campdera.
Eat. =
Amos Eaton.
Cand.
DeCandolle, usually DC.
D. C. Eat.
D. C. Eaton, grandson.
Casp.
Caspary.
Edgew.
Edgeworth.
Cass.
Cassini.
Edw.
Edwards.
Catesb.
Catesby.
Ehren.
Ehrenberg.
Cav.
Cavanilles.
Ehrh.
Ehrhart.
Cerv.
Cervantes.
Eicfil.
Eichler.
Cham.
Charaisso.
Eiseng.
Eisengrein.
Chapm.
A. W. Chapman.
Ell.
Elliott.
Chav.
Chavannes.
EndL
Endlicher.
Cltois.
'Choisy.
Engelm.
Engelmann.
Clayt.
Clayton.
Enyl.
Engler.
Clus.
Clusius.
Eschs.
Eschscholtz.
Collad.
Colladon.
Eschw.
Eschweiler.
Colm.
Colmeiro.
Ettingsh.
Ettingshausen.
Comrn.
Commelin.
Corn.
Cornuti.
Fendl.
Fendler.
Coss.
Cosson.
Feuil.
Feuillee.
Cunn.
Cunningham, A. or J.
Finger h.
Fingerhuth.
Curt.
Win. Curtis.
Fisch.
Fischer.
M.A.Curt. M.A.Curtis.
Forsk.
Forskai.
Forst.
Forster.
Dalech.
Dalechamps.
Foam.
Fournier.
Dalib.
Dalibard.
Fresen.
Fresenius.
Darl.
Darlington.
Freyc.
Freycinet.
DC.
1
Fred.
Froelich.
DeC.
> A. P. DeCandolle.
A. DC.
Alphonse DeCandolle,son.
Gcertn.
J. Gasrtner.
Cos. DC.
Casimir DeCandolle, the
Gcertn.f.
C. T. Gaertner.
Decne.
Decaisne. [grandson.
Gardn.
Gardner.
Deless.
Delessert.
Garid.
Garidel.
Dennst.
Dennstedt.
Gasp.
Gasparrini.
Desc.
Deseourtilz.
Gaud.
Gaudin.
Desf.
Desfontaines.
Gaudich.
Gaudichaud.
Desj.
Desjardins.
Germ.
Germain.
Desmar.
Desmazieres.
Gesn.
Gesner.
Desmoul.
Desmoulins.
Gilib.
Gilibert.
Desv.
Desvaux.
Ging.
Gingins de Lassaraz.
Dicks.
Dickson.
Gis.
Giseke.
Diesb.
Diesbach.
Gled.
Gleditsch.
Dieter.
Dieterich.
Gleich.
Gleichen.
Dietr.
Dietrich.
Glox.
Gloxin.
Dili
Dillenius.
Gmel.
J. G. Gmelin.
Dillw.
Dillwyn.
C. Gmel.
C. C. Gmelin of Baden.
Dod.
Dodonseus (Dodoens).
S. Gmel.
S. G. Gmelin.
D'Orb.
D'Orbigny.
Godr.
Godron.
Dorst.
Dorstenius.
Gcepp.
Goeppert.
Dougl.
Douglas.
Good.
Goodenough.
Drej.
Drejer.
Gren.
Grenier.
Drt/and.
Dryander.
Grev.
Greville.
Dufr.
Dufresne.
Griseb.
Grisebach.
Duham.
Duhamel du Monceau.
Green.
Greenland.
Dumort.
Dumortier.
Gron. )
Gronovius.
Dun.
Dunal.
Gronov. )
ABBREVIATIONS.
387
Guett. =
Guettard.
Jacq. =
N. J. Jaequin.
Gnib.
Guibord.
Jacq.f.
J. F. Jaequin, son.
Guil/em.
Guillemin.
J. St. Hil.
Jaume !St. Hilaire.
Gaimp.
Guimpel.
Jord.
Jordan.
Gunn.
Gunnerus.
Junyh.
Junghulm.
Guss.
Gussone.
Juss.
A. L. Jussieu.
Adr. Juss.
Adrien Jussieu, son.
Hagenb.
Hagenbach.
Hail.
Ham.
Haller.
Hamilton.
Kcemp.
J^ctrst,
Kaempfer.
Karsten.
llanb.
Hanst.
Hartm.
Hartw.
Hanbury.
Hanstein.
Hartmann.
Hartweg.
Kaulf.
Kindb.
Kirschl.
Kit.
Kaulfuss.
Kindberg.
Kirschleger.
Kitaibel.
Haru.
Harvey.
Kcelr.
Koelreuter.
lias*.
Hassk.
Hausm.
Ha iv.
Helens.
Hedw.
Hassall.
Hasskarl.
Hausmann.
Ha worth.
Hebenstreit.
Hedwig.
Korth.
Kostel.
Kremp.
Kromb.
Kuetz.
Korthals.
Kosteletzky.
Krempelhuber.
Krombholz.
Kuetzing.
JJegelm.
Hegelmaier.
Hegetsch.
Hegetschweiler.
L.
Linnaeus.
Heist.
Heister.
Labill.
La Billardiere
Heldr.
Heldreich.
Last.
La3stadius.
Helw.
Helwing.
Lag.
Lagasca.
Hemsl.
Hemsley.
Lall.
Lallement. [Marck).
Henck.
Henckel.
Lam.
Lamarck (Monnet de La
Henfr.
Henfrey.
Lamb.
Lambert.
Hensl.
Henslow.
Lamour.
Lamouroux.
Herb.
Herbert.
Langsd.
Langsdorf.
Herm.
Hermann.
La Peyr.
La Peyrouse.
mid.
Hildebrand.
La Pyl.
La Pylaie.
Hochst.
Hochstetter.
Ledeb.
Ledebour.
II off m.
G. F. Hoffmann.
Lehm.
Lehmann.
H. Hoffm
Hermann Hoffmann
Lem.
Lemaire.
Hoffmanns. Hoff mannsegg.
I^esq.
Lesquereux.
Hofm.
Hofmeister
Less.
Lessing.
Hohen.
Hohenacker.
Lestib.
Lestiboudois.
Holmsk.
Holmskiold.
Lo.
Leveille'.
Jiomb.
Hombron.
L'Hcr.
L'Heritier.
Hook.
Win. J. Hooker.
L'Herm.
L'Herminier.
Hook.f.
J. D. Hooker, son.
Liebm.
Liebmann.
Hopk.
Hopkirk.
Light/.
Lightfoot.
Hornem.
Hornemann.
Lilij.
Lilijeblad.
Hornsch.
Hornschuch.
Lindb.
Lindberg.
Horsf.
Horsficld.
Lindbl.
Lindblom.
Houst.
Houston.
Lindenb.
Lindenberg.
Houtt.
Houttuyn.
Lindh.
Lindheimer.
Huds.
Hudson.
Lindl.
Lindley.
Hueb.
Huebener.
Linn.
Linnaeus. Also L.
Humb.
Humboldt.
Linn.f.
C. Linnaeus, son.
HBK.
( Humboldt, Bonpland, and
Lodd.
Loddiges.
( Kunth.
Lffjl.
Lcefling.
388
ABBREVIATIONS.
Loes. == Loeselius.
Naud. =
Naudin.
Lois. Loiseleur-Delongschamps.
Neck.
Necker.
Loud. Loudon.
Lour. Loureiro.
Nees or )
N.abE. }
C. F. Nees von Esenbeck.
Lid iff. Ludwig.
T. Nees
T.F.L. Nees von Esenbeck.
Luinn. Lumnitzer.
Nestl.
Nestler.
Lyngb. Lyngbye.
Newb.
Newberry.
Newm.
Newman.
Macf. Macfadyen.
Ncegg.
Noeggerath.
Macgil. MacGillivray.
Nois.
Noisette.
Magn. Magnol. [stein.
Nord.
Nordstedt.
M. Bleb. Marschall von Bieber-
Not.
Notaris.
Marsh. Humphrey Marshall.
Nutt.
Nuttall.
Mars. Marsili.
Nyl.
Nylander.
Mart. Martius.
Nym.
Nyman.
Mass. Massalongo.
Mast. Masters.
(Ed.
OEder.
Maxim. Maximowicz.
(Erst.
CErsted.
Med. Medikus or Medicus.
Oliv.
Olivier.
Meisn. \ Meisner or Meissner.
Meissn. }
D. Oliv.
Orb.
D. Oliver.
A. or C. d'Orbigny.
Meneg. Meneghini.
Orph.
Orphanides.
Menz. Menzies.
Ort.
Ortega.
Mert. Mertens.
Oudem.
Oudemans.
Metten. Mettenius.
Mich. Micheli.
P.deBeauv.Psdisot de Beauvois.
Michx. ) Andn < Michaux
Pall.
Pallas.
MX. )
Panz.
Panzer.
Michx. f. F. A. Michaux, son.
Park.
Parkinson.
Midden. Middendorff.
Pad.
Parlatore.
Mill. Philip Miller.
Pasq.
Pasquale.
Mill. J. John S. Mueller or Miller.
Pa,-.
Pavon.
Miq. Miquel.
Perl.
Perleb.
Mirb. Mirbel.
Pers.
Persoon.
Mitch. John Mitchell.
Phiiib.
Philibert.
Mitt. Mitten.
Planch.
J. E. Planchon.
Moc. Mo9ino.
G. Planch
Gustave Planchon.
Molk. Molkenboer.
Pluk.
Plukenet.
Mont. C. Montagne.
Plum.
Plumier, Lat. Plumerius.
Moq. Moquin-Tandon.
Pcepp.
Poeppig.
Moric. Moricand.
Poir.
Poiret.
Moris. Morison.
Poit.
Poiteau.
Morr. Morren.
Poll.
Pollich.
Moug. Mougcot.
Post.
Postels.
Muell. Arg. J. Mueller of Argau.
Pourr.
Pourret.
F. Muell. Ferdinand Mueller.
Pringsh.
Pringsheim.
0. Muelt. Otto Mueller of Denmark.
Pritz.
Pritzel.
MM. Muhlenberg.
Putter.
Putterlich.
Munt Munting.
Murr. J A. Murray.
Rdbenh.
Rabenhorst.
A. Murr Andrew Murray.
Radlk.
Radlkofer.
Raf.
Rafinesque-Schmaltz.
Nacc. Naccari.
Rasp.
Raspail.
Ncey. Naegeli.
Red.
Redoute.
ABBREVIATIONS.
389
Reich. = Reichard.
Scop. = Scopoli.
Reichenb. H. G. L. Reichenbach.
Seem.
Seemann.
Reichenb.f. H. G. Reichenbach, son.
Sendt.
Sendtner.
Reinw. Reinwardt.
Seneb.
Senebier.
Reiss. Reisseck.
Ser.
Seringe.
Retz. Retzius.
Seub.
Seubert.
Reut. Reuter.
Sibth.
Sibthorp.
Rich. L. C. Richard.
Sieb.
Sieber.
Rich./, i Achille Richard.
A. Rich. }
Sieb.
Soland.
Siebold.
Solander.
Richards. John Richardson.
Sow.
Sowerby.
Richt. Richter.
Spenn.
Spenner.
Ridd. Riddell.
Spreng.
Sprengel.
Riu. Rivinus.
Sternb,
Sternberg.
Rop.hl. Roehling.
Steud.
Steudel.
Rcem. J. J. Roemer.
Stev.
Steven.
M. J. Roem.M. J. Roemer.
Sull.
Sullivant.
^ m - & 1 Roemer & Schultes.
oc/i. )
Sw.
Swartz.
Roep. Roeper.
Targ.
Targioni-Tozetti.
Rohrb. Rohrbach.
Ten.
Tenore.
Rostk. Rostkovius.
Thorns.
Thomas Thomson.
Rothr. Rothrock.
Thuill.
Thuillier.
Rottb. Rottbcell.
Thunb.
Thunberg.
Rottl. Rottler.
Thurb.
Thurber.
Roum. Roumegere.
Thurm.
Thurman.
Roxb. Roxburgh.
Tod.
Todaro.
Roy. Royen.
Ton:
Torrey.
Rudb. Rudbeck.
Torr.fr
Gr.Torrey & A. Gray.
Rupr. Ruprecht.
Tourn.
Tournefort.
Trait.
Trattinick.
Sacc. Saccardo.
Traut.
Trautvetter.
Sadl. Sadler.
Trev.
Treviranus.
St. Hil. A. Saint-Hilaire.
Trin.
Trinius.
Salisb. Salisbury.
Tuck.
Tuckerman.
Salm-Dyck.Prince Jos. Salm-Riffer-
Turcz.
Turczaninow.
Sauss. Saussure. [schied-Dyck.
Turn.
Turner.
Schimp. Schimper.
Turp.
Turpin.
Schk. Schkuhr.
SMecht. Schlechtendal.
Vaill.
Vaillant.
Schleich. Schleicher.
VeiU.
Veillard or Vieillard.
Schomb. Schomburgh.
Vauch.
Vaucher.
Schrad. Schrader.
Vent.
Ventenat.
Schreb. Schreber.
Vill.
Villars, or Villar.
Schneb. Schuebeler.
Vis.
Visiani.
Srhult. Schultes.
Vittad.
Vittadini.
Schultz 1 C. H. Schultz, Bipontinus
Viv.
Viviani.
Bip. ) (Zweibrucken)
Vog.
T. Vogel.
Schum. Schumacher.
Schnitzl. Schnitzlein.
Wahl.
Wahlenberg.
Schwcegr. Schwaegrichea
Wahlst.
Wahlstedt.
Schwein. Schweinitz.
Walds.
Waldstein-
Schwei.if. Schweinfurth.
Wall.
Wallich.
Schwend. Schwendener.
Wallm.
Wallman.
390
ABBREVIATIONS.
Wattr. =
Wdp.
Walt.
Wang.
Warm.
Wats.
H.C.Wats.
S. Wats.
Web.
Wedd.
Weinm,
Welw.
Wender.
Wendl.
Wiks.
Wallroth.
Walpers.
Walter.
Wangenheim.
Warming.
P. W. Watson.
H. C. Watson.
Sereno Watson.
Weber.
Weddell.
Weinmann.
Welwitsch.
Wenderoth.
Wendland.
Wikstrom.
Wildb. =
Wildbrand.
WiUd.
Willdenow.
Willk.
Willkomm.
Wils.
Wilson.
Wimm.
Wimmer.
Wisliz.
Wislizenus.
With.
Withering.
Woodv.
Woodville.
Wulf.
Wulfen.
Zanard.
Zanardini.
Zetterst.
Zetterstedt.
Zu.cc.
Zuccarini.
Zuccag.
Zuccagini.
2. ABBREVIATIONS OF NAMES OF ORGANS AND TERMS
USED IN BOTANICAL WRITINGS.
jEst. ^Estate, in summer.
sst. ^Estivation.
Alb. Albumen.
Anth. Anther.
Art. Artificial.
Auct., Auctt. Auctorum, of authors.
Aut. Autumnal.
B. or Beat. Beatus, " the late," re-
cently deceased.
Br. Bract.
Cal. Calyx.
Cel. Celeberrimus, or Very cele-
brated.
Cent. Centimetre.
Cl Clarissimus.
Char. Character.
Coll. Collection.
Car. Corolla.
Cult. Cultivated.
Decim. or Dec. Decimetre.
Descr. Description
Dlff. Differentiae, the distinguishing
marks.
Ed. Edition.
Embr. Embryo.
Ess. Essential, as Char. Ess.
Excl. Excluding, or being excluded.
Excl. Syn. Excluding the synonym
or synonyms.
Fam. Family.
Fil. Filament of the stamen.
Fl. Flower (flos); Flora, or some-
times Floret, it flowers.
Fcem. Female plant, flower, &c.
Fol. Folium, leaf.
Fr. Fruit.
Fructif. Fructification.
Gen. Genus or Generic.
Germ. Germen, Linnsean name for
ovary ; also Germination.
H. Herbarium.
Hab. Habitat, place of growth ;
sometimes for Habeo, I have.
Herb. Herbarium.
Hort. Hortus, garden.
Hortul. Hortulanorum, of the gar-
deners.
7c. Icon, a plate or figure.
///. Illustris, illustrious.
Ined. Unpublished.
Inf. Inferior.
In/I. Inflorescence.
Inv. Involucre.
Lat. Lateral, or relating to width.
Lin. Linea, a line (the 12th of an
inch).
Lit., Litt. In a letter or letters.
/. c. Loco citato, in the place cited.
ABBREVIATIONS.
891
M isc. Male plant, flower, &c.
Mill, or mm. Millimetre.
Mss. Manuscripts.
Mus. Museum.
N. or No. Number.
Nat. Natural.
Norn. Nomen, name.
Obs. Observation.
Ord. Order.
Oo. Ovary.
/>. Page, or sometimes Part.
Fed. Peduncle or Pedicel, or
Pedalis, a foot long or high.
Ptric. Pericarp.
Perig. Perigonium.
Pet. Petal or Petiole.
Pist. Pistil.
Plac. Placenta.
Poll. Pollicaris, an inch long.
p. p. Pro parte, in part.
Prodr. or Prod. Prodromus.
Rud. Radix, root ; or Radical.
Ram. Ramus, branch.
s. Seu, or Sive, Latin for or.
Sect.. Section.
Segm. Segment.
Sem. Semen, seed.
for
Sep. Sepal.
Ser. Series.
Sice. Siccatus or Siccus, dried or dry.
Spec, or Sp. Species, or specimen.
Spont. Spontaneous.
Stain. Stamen or Staminate.
Sup. Superior.
Syn. Synonym or Synopsis.
T. or Tab. Tabula, plate.
T. Tomus, volume.
V. Volume : sometimes for Vel, or;
sometimes Vide, see.
Far. Variety.
Veg. Vegetation, characters of
Fern. Vernal.
v. s. Visa sicca, or Vidi siccam.
v. v. Visa viva, or Vidi vivam; the
first indicating that a dried speci-
men of the plant, the second that
the living plant has been exam-
ined.
v. s. c. and v. s. s., indicates that
the dried specimen was cultivated
(c) or spontaneous (s).
v. v. c. and v. v. s., that the living
plant seen was cultivated (c) or
spontaneous (s).
SIGNS.
1. SIGNS USED BY LINNAEUS.
Q An annual plant.
$ A biennial.
2/ A perennial,
b A tree or shrub.
# Affixed to a reference, means that a good description will be found there,
f Indicates an obscure or doubtful species.
2. SIGNS USED BY DECANDOLLE AND LATER
WRITERS.
O A monocarpic plant, i. e. which dies after once flowering and fruiting,
either annual or biennial, or of longer duration.
QU Annual.
Biennial.
@ Monocarpic perennial, such as Agave.
392 SIGNS.
^ Perennial herb.
^ Suffrutex, an undershrub.
*> Frutex, a shrub.
5 Arbuscula, a tree-like shrub of ten to twenty -five feet in height.
5 Arbor, a tree.
r\ A climbing plant.
A An evergreen.
$ Male plant or flower.
9 Female .plant or flower.
^T Hermaphrodite plant or flower.
co Indefinitely numerous, e. a. coandra, polyandrous.
? A sign of doubt. " Thalictrum ? Japonicum," doubts if the plant is
really a Thalictrum. " Thalictrum Japonicum, Thunb.? " doubts if the
plant in hand is truly the species of Thunberg. Thalictrum Japoni-
cum, Thunb., Willd. 1 ? doubts whether Willdenow's T, Japonicum is
really that of Thunberg.
! A sign of certainty. As " Thalictrum anemonoides, Michx. ! Fl. Bor. Am.
p. 322," as used by DeCandolle, affirms that he has seen an authentic
original specimen of this author. Affixed to the name of a collector,
as " Virginia, Clayton ! " it affirms that the writer has examined a
specimen collected by the person to whose name it is appended.
- Between two figures, as in " Stamens 5-10," indicates the extremes of
difference, as that the stamens are from five to ten.
' " The signs for degrees, minutes, and seconds, as 1, 2', 3", are used in
Gray's Manual of Botany of the Northern United States, for feet (),
inches ('), and lines ("). With European authors, usually the sign
for minutes is for feet ; that of seconds for inches : thus 1', a foot
high ; I", an inch long ; and I"', a line long.
O= Cotyledons accumbent to the radicle.
OH Cotyledons incumbent on the radicle.
GLOSSARY,
OR
DICTIONARY OF BOTANICAL TERMS, ENGLISH
AND LATIN,
COMBINED WITH AN INDEX.
THIS Glossary is intended to contain all the principal technical terms (substan-
tive as well as adjective) of structural and systematic Botany, as far at least as
concerns Phaenogamous plants. Most of the special terms relating to the lower
CryptogamSa and to Vegetable Anatomy and Physiology are relegated to the vol-
umes devoted to those departments. The annexed numbers refer to pages of this
volume. Very many of the terms are seldom employed, or are wholly out of use.
The principal Latin terms are given separately only when there is no English equiv-
alent differing merely in the termination. When the word is essentially the same,
the Latin termination (of adjectives in the nominative masculine only) is annexed
to the English word in a parenthesis. The changed termination goes back mostly
to the penultimate consonant. It is unnecessary in a work like this to accentuate
all the technical words; but, in the case of words liable to mispronunciation, an
accent-mark is placed over the syllable which takes the principal accent. The
glossary, as here drawn up, may serve to indicate the meaning of the commoner
descriptive specific names of plants.
A, privative, as the initial in many
words of Greek derivation, signifies
the absence of the organ mentioned;
as, ylpetalous, destitute of petals;
Aphyllous, leafless. In words be-
ginning with a vowel, this prefix is
changed to an; as, ylnanthous, flow-
erless ; ^nantherous, antherless.
Abbreviations, 385.
Aberrant. Wandering, applied to spe-
cies, genera, &c., which differ in some
respect from the usual or normal char-
acter of the group they belong to.
Abnormal (Abnormis). Differing from
the normal or usual structure.
Aboriginal. Strictly native; indigenous.
Abortion (Abortus). Imperfect develop-
ment or non-development of an organ ;
179, 187.
Abortive (-ivus). Defective or barren.
Abrupt (Abruptus). Terminating sud-
denly ; the opposite of tapering.
Abruptly pinnate. Pinnate without a
terminal leaflet or appendage ; 101.
Acanthocladous (-us). Having spinal
branches.
Acantkophorous (-us). Spine-bearing.
Acaulescent (-ens). Stemless, or appar-
ently so, with no proper caulis ; 45.
Acaulis. Stemless ; same as Acaulescent.
Accessory. Something additional, or of
the nature of appendage.
Accessory Buds, 44.
Accessory Fruits, 300.
Accrescent (-ens). Increasing in size
with age, as often occurs with the
calyx after flowering.
Accrete (-us). Grown together, or con-
solidated with some contiguous body.
Accumbent (-ens). Lying against an-
other body.
Accumbent Cotyledons. With
against the radicle; 313.
Acephalous (-us). Headless.
394
GLOSSARY.
Acerose (-6sus). Needle-shaped, like the
leaves of Pines.
Acetdbuliform (-ormis). In the form of
a shallow open cup or saucer.
AchoBUium or Achenium. A small, dry
and hard, one-celled, one-seeded, inde-
hiscent fruit ; strictly one of a single
and free carpel ; but extended to simi-
lar ones of more than one carpel, and
also with adnate calyx; 294. (Achce-
nium is etymologically the proper
orthography ; but achenium is be-
coming the commoner form.)
Achcenocarp (-arpium). General name
of a dry and indehiscent fruit ; 292.
Achf.nodium. Such a double achenium
as that of Umbelliferae ; a Cremo-
carp.
Achlamydeous (-eus). Destitute of peri-
anth; 191.
Acicula. A bristle.
Acicular (-am 1 ). Bristle-shaped, or slen-
der needle-shaped.
Acindctform (-ormis). Scymitar-shaped ;
curved with rounded point, thicker on
the straighter edge than on the con-
vex edge.
Acinosus. Like grapes or grape-seed.
Acinus. Classically a berry, particu-
larly a grape, or its stony seed, or' a
bunch of berries; now sometimes ap-
plied to the separate carpels of an
aggregate baccate fruit, or to the con-
tained stone or seed ; 297.
Acorn. Fruit of the Oak.
Acotyledon } p\. Acotyledons, Acotyledones.
A plant or plants destitute of coty-
ledon, or
Acotyledonous (-eus). Without cotyle-
dons ; as the embryo of Cuscuta ; 26,
38. Mostly applied, as bv Jussieu,
to plants which have no proper seed
nor embryo, and therefore no co'yle-
don ; 339.
Acramphibrya. Plants producing side as
well as terminal buds or growths ; 341
Acrobryn. Plants growing from apex
^only; 341.
Acroyen (Acrdgence). Name of class of
plants which in growth are said to be
Acrof/enous. Growing from the ap<>x
or by terminal buds only.
Acrosdrcum . Desvaux's name for a
berry from an ovary with adnate
calyx.
Acro.opira. An old name of the plu-
mule of a grain in germination.
Aculeate (-eatus) Prickly; beset with
aculei.
Aculeosus. Abounding with prickles.
Aculeolate (-atus). Beset with diminu-
tive prickles, or
Aculeoli. Diminutive of aculei.
Aculeus. A prickle ; a pointed small
excrescence of the bark.
Acumen. A tapering point.
Acuminate (-atus). Ending in a tapering
point ; 96.
Acute (Acutus). Terminating in an acute
angle; 97.
Acropetal. Developing from below up-
ward, or from base toward apex.
Actinomorphous (-us). Capable of bi-
section through two or more planes
into similar halves, as is a regular
symmetrical flower; 175.
Acutiusculus. Somewhat acute ; acutish.
Adelpkous (-us, Adelphi, brothers). Sta-
mens with coalescent or clustered fila-
ments are monadelphous, diadelphous,
&c., according to the number of Adtl-
pliia or brotherhoods.
Aden. Greek for gland, is compounded
with Greek words with this meaning;
as, Adenophorus, gland-bearing; Ad~
enopliyllus, leaves bearing glands, &e.
Adtjlutinate (-atus). Same as accrete
Adherent (Adhcerens). Generally same
as adnate ; may refer to adhesions not
congenital.
Adnate (-atus). Congenitally united to ;
as the calyx-tube of the gooseberry to
the ovary; 182. Adnate ant I. er is one
seemingly borne on the outer or inner
face of the filament ; i. e. extrorsely or
introrsely fixed by its whole length to
the connective ; 253.
Adnatwn. The state of being adnate;
179, 181.
Adpressus Latin of appressed.
Adscendens. Latin of ascending.
Adsuryens. Latin of assurgent.
Adventitious, Adrentice. That which has
come from abroad or as a stranger ; as
a plant lately or by chance introduced
from another country.
Adventitious Buds; 45.
jEqmlnterctUs. Equilateral, equal-sided.
yEqiKilifloi'us. When all the flowers of
the same head or cluster are alike in
form as well as character.
jfi'iudlis, jEqunns. Equal ; equalling.
Aerial roots, &c. ; 33.
Aerophytes. Air-plants; 35.
^Eruyinosus. Verdigris-colored.
JEstival (-dlis). Relating to summer.
^Estivation (-io). The disposition of the
parts of a flower in the bud ; 132.
GLOSSARY.
395
jEterio. A form of aggregate fruit ; 300.
sEtheogamia, ^Ethoyamous, 340.
Affinity. True and near relationship;
327, 330.
Ayamous or Agamic. Destitute of sexes.
, ( Heaped or crowded
Agglomerate. \ .
. , . > < into a dense cluster,
Aggregate (-atus i. )
v. but not cohering.
Aggregate Fruits. Those formed of
aggregate carpels of the same flower ;
298, 301.
Agrestis. Growing in fields.
Air-plants. Plants unconnected with the
ground; 35.
Akene, Akenium. See Achaenium.
Ala (pi. alee). A wing. Also the side
petals of a papilionaceous corolla;
185. Has also been used in the sense
of axilla.
Alabdstrum. A flower-bud; 40.
Alar (Alaris). From ala in the sense of
axilla, therefore axillary or in the forks.
Alnte (-atus). Winged.
Albescent!, Albicans. Whitened, whitish,
or hoary.
Albumen of the seed. Any deposit of
nutritive material within the seed-
ci ats, and not in the embryo ; 14, 309.
Albuminous or Albuminose (Albumincsuf).
Said of seeds provided with albumen ;
13, 309.
Alburnum. Sapwo d: the newer wood
of an exogenous stem ; 80.
Albus. White.
Allagostemonous. With stamens alterna-
tively inserted on the torus and on the
petals.
Alliaceous (-eus). Having the smell of
garlic.
Alliance. Synonym of Cohort ; 320.
Allogamy. Fecundation of the ovules of
a flower by other than its own pollen ;
cross fertilization, 216.
Alj> strine (Alpestris). Growing on
mountains below an alpine region
or one unwoodtd from cold.
Alpine (-inus). Growing on the higher
parts of the Alps, or (by extension of
meaning) on other mountains above
the limits of trees.
Alternate (Altf'rnus). One after an-
other; as rf leaves placed singly
instead of in pairs (opposite) or in
whorls. Also, standing before inter-
vals ; as stamens alternate with petals
instead of before them; 6, 119.
Alternative (-tews). In aestivation, with
an inner whorl alternating with an
outer one; 134, 13(j.
Alveolate(-atus). Honeycombed; having
deep angular cavities (Alveoli) sepa-
rated by thin partitions, as the recep-
tacle of cotton-thistle.
Jlmbitus. The ray or circumference of
a head, &c.
Ament (Amentum). A catkin, or pe-
culiar scaly spike ; 150.
Amentaceous (-eus). Bearing catkins, or
cat kin- like.
Amorphous (-us). Shapeless; of in-
definite form.
Amphanthium. One of the (needless)
names coined for a dilated receptacle
of inflorescence.
Amphibrya. Equivalent to Monocotyle-
dones; 341.
AmpJi icarpous (-us). Producing two
kinds of fruit.
Ampldyamous Cryptogams, 340.
Amj>Jtiynstri(t. Peculiar leaves (of He-
paticte) imitating stipules.
Amphisarca. A hard-rinded berry, or
fruit succulent within, and woody or
crustaceous without, as a calibash.
Ampliispermium. Link's name for a
one-seeded pericarp which is con-
formed to the seed ; an akene.
Amphitropous (-us), wrongly Amphi-
tropal. Turned both ways; applied
to an ovule with hilum intermediate
between micropyle and chalaza; 279.
Jlmphora. A pitcher; and the lower
part of a pyxis.
Ampltctens, Amplexans, Amplexus Em-
bracing, clasping.
Amplexicaul (-aulis). Clasping a stem,
as does the base of certain leaves.
Ampliate (-atus). Enlarged or dilated.
Ampulla. A bladder or flask-shaped
organ, as of Utricularia.
Ampullaceous (-us), or Ampullatform.
In the form of a bladder or short flask.
Amylaceous (-eus). Resembling or com-
posed of starch, or Amylum.
Amyloid. Analogous to starch.
Analogy (Analoyia). Likeness in cer-
tain respects. As distinguished from
affinity, it means resemblance in cer-
tain respects only, not in the plan of
structure. Thus, a Ranunculus is
analogous to a Potentilla, but there
is no near affinity or relationship be-
tween the two. And the tendril of a
Pea, that of a Smilax, and that of
the Grape-vine are analogues ; i. e , are
analogous organs, but are not homo-
lofjues ; for the first answers to a leaf,
the seeped tq stipules, and the third
396
GLOSSARY.
to a stem. The spur of a Larkspur is
analogous to one of the tive spurs of
Columbine, but not homologous with
it; for the first is a sepal, and the
second a petal.
Anandrous. Destitute of stamens.
Anantherous (-us). Destitute of anthers.
Ananthous (-us). Flowerless.
Anastomosis- The connection of veins,
&c., by cross-veins, forming reticu-
lation.
Andtropous (-us), wrongly Anatropal.
The reversed ovule, with micropyle
close bv the side of the hilum, and
chalaza at the opposite end ; 279.
Anceps, Engl. Ancipital. Botanically
always used in the sense of two-
edged.
Ander, andra, andrum. In Greek com-
pounds, the male.
Andro-ditxcious. With flowers on one
plant hermaphrodite, and on another
staminate only; 191.
Andr&cium. The stamens of a flower
collectively ; 165, 249.
Androgynous (-us). Said of an inflores-
cence composed of both male and
female flowers.
Aiulrophoi-e (Andrtphorum). A sup-
port or column on which stamens are
raised.
Androus. See Ander.
Anemophilous. Literally wind-loving.
Said of flowers which are fecundated
by wind-borne pollen; 217.
Anfractuosus. Abruptly bent hither and
thither, as the stamens of Cucur-
bita.
Anyiocarpous (-us). When a fiuit is
covered by some envelope.
Anyiospermia. A Linna-an artificial
order; 337.
Anyiospermous, Anyiospermce, Anyio-
sperms. Plants with seeds borne in a
pericarp; 259.
'Angular Divergence of leaves ; 123.
Anisomerous (-us). Unequal in number
in the different circles of the flower;
unsymmetrical.
Anisope.talous (-us). With unequal petals.
Anisophyllous (-us). Unequal-leaved;
i. e., the two leaves of a pair unequal.
Anisostemonous (-us). When the sta-
mens are not of the number of the
petals.
Annotinus. A year old, or in yearly
growths.
Annual (Annuus). Of only one year's
duration; 30.
Annular (-aris). In the form of a ring;
or marked transversely by rings.
The latter more properly.
Annulate (-atus). Marked with rings.
Annulus. A ring, such as that with
which the sporangia of some Ferns
and Mosses are furnished.
Anophytes (Anophyta). Name of group
comprising Mosses, &c.
Anttposition Same as Superposition;
179, 195.
Anterior, as to position, denotes the front
side, or averse from the axis of inflo-
rescence; 160.
Anthela. A deliquescent and paniculate
cyme, with median ramification, and
the lateral axes overtopping the
central, as in Juncus tenuis, &c.
May be either a Drepanium or a Khi-
pidium.
Anthemy, Anthemia. A flower-cluster
of any kind ; 144.
Anther (Anthera). The polliniferous
part of a stamen; 165, 251.
Antheridium. An analogue of the an-
ther in Cryptogams.
Anthtriferous (-us). Anther-bearing.
Jlnthtsis. The time at which a flower
is perfected and opens; or the act of
expansion of a flower.
Anthocarpous(-us)i Anthocarpium. Fruits
in which some organ exterior to the
pericarp is concerned ; 300.
Anthoclinium. Name of a receptacle of
inflorescence, such as that of Com-
positse.
Anthodium. A name for the head of
flowers (or so-called compound flowers)
of Composite; 147.
Antholysis. A retrograde metamor-
phosis of a flower, in which normally
combined parts are separated.
Anthophore (Anthophorum). The stipe
when developed between calyx and
corolla; 212.
Anthus or Anthos. A flower, in Greek
compounds.
Anticous (Anticus). Facing anteriorly;
253.
Antitropous (-us), less properly Antitro-
pal. Said of an embryo with radicle
pointing to the end of the seed oppo-
site the hilum; 312.
Antrorse, Direpted upward or forward.
Apetalous (-#). Having no petals; 190.
Ape$. Besides its ordinary meaning,
the top of a thing, it was once the
technical name of an anther-; 166,.
AphylLus (-us). Leafless.
GLOSSARY.
397
Apical (-alls). Relating to the apex or tip.
Apices. The name for anthers anterior
to Ludwig and Linnaeus; 166.
Apiculate (-us). Ending in a short
pointed tip or apicula.
Apocarpous (-us). When carpels of a
gynoicium are separate; 261, 262.
Apophysis. An enlargement or swelling
of the surface of an organ at some par-
ticular part.
Apothecia. The "shields " or fructify-
ing disks of Lichenes.
Apotropous (-us). Said of an anatropous
ovule which when pendulous has
rhaphe averse ; 282.
Appendage, Appendix. Any superadded
or subsidiary part.
Appendiculate (-atus). Furnished with a
small appendage (Appendiculum), or
with any appendage.
Appositus. Placed side by side.
Appressed (Lat. Adpressus). Lying flat
against or together for the whole
^length.
jlpricus. Growing in diy sunny places.
Apterous (-us). Wingless; not alate.
Aquatic (-icus). Living in water.
Aquatilis. Living under water.
Arachnoid (-oideua). Cobwebby ; com-
posed of slender entangled hairs.
Araneose (-osus), Araneus. Like spider-
web; same as Arachnoid.
Arbor. A tree ; 50.
Arboreous (Arboreus). Tree-like, or re-
lating to a tree,
Arborescent (-ens). Tree-like ; approach-
ing the size of a tree.
Arboretum, also Arbustum. A place
where trees are grown ; an arranged
collection of trees.
Arbuscula. A small shrub of tree-like
growth or form.
Arbuscularis. Ramified like a little tree.
Archegonium. The spore-case of mosses,
&c., in an early state.
Arcuate (-atus). Moderately curved, as
if bent like a bow.
Areola, pi. Areolce. Spaces marked out
on a surface, as by the reticulation of
veins, &c.
Areolate (-atus). Marked with areolae.
Arenosus, Arenarius. Growing in sand
or sandy places.
Argentate (-atus, Arg*>nteus). Silver}',
or shining white with a tinge of gray.
Aryillosus. Growing in clayey soil.
Aryos. Greek for pure white, used in
compounds ; as, aryophyllus, white-
leaved.
Argutus. Sharp-toothed; said of the
serration of leaves.
Aryyros. Greek for silvery ; used in
compounds; as,argyrophyllus, silvery-
leaved.
Arhizal (Arhizus). Rootless.
Arillatt (-atus). Having an arillus.
Aril, Arillus. An extraneous or late-
formed seed-coat or covering, or an
appendage growing from or about the
hi linn of a seed ; 308. (
Arilli/'vrm (-ormis). In the form of an,
arillus.
Arillode, Arillodium. A false arillus,
or one which does not originate from
or below the hilum, but from the
mkropyle or rhaphe; 309.
Arista. An awn.
Aiistate (-atus). Awned; bearing an
arista.
Aristulate (-atus). Bearing a diminu-
tive awn.
Arrect (Arrectus). Brought into an
upright position.
Arrow-shaped, Arrow-headed. Same as
Sagittate; 96.
Articulated (-atus). Jointed, or having
the appearance of a joint or articula-
tion (Articulus). As of the word joint
itself, the context must show whether
the articulations mean the portions
which are connected by a joint, or the
place of connection.
Artificial Classification, 331.
Ascending ( Adscendens). Rising upward.
Sometimes used for directed upward,
as when the stem is termed the As-
cending Axis (11); more commonly
denotes curving or rising obliquely
upward; 53.
Ascidium. A pitcher-shaped or flask-
shaped organ or appendage; 111.
Ascus. A sac ; a kind of spore-cases, as
in certain Fungi and Lichenes
Aspdragi. A name for Turiones, or any
scaly shoots from underground, as
those of Asparagus.
Aspergilliform (-ormis). Brush-shaped,
i. e. like the aspergillum, or brush
used to sprinkle holy water ; made up
of numerous spreading hairs, &c., in
a tuft, as the stigmas of Grasses.
Asperous (Asper). Rough to the touch.
Assimilation. The action or process by
which extraneous matter or crude food
is converted into vegetable matter.
Assuryent (Adsurgens). Rising or curv-
ing upward : 53.
Astichous (-us). Not in rows.
398
GLOSSARY.
Astomous (-us). Without a stoma or
mouth.
Atavism (-mm). Ancestral resemblance.
Ater. Pure black.
jftkera. Greek for Arista or Awn.
At> atus. Blackened or turning black.
Mrvpous (-us), wrongly Atropal. Not
turned ; applied to an ovule the same
as orthotropous ; 277.
Attenuate (-atus). Slenderly tapering
or narrow.
Auctus. Same as accrescent; enlarged
after flowering ; augmented by an ad-
dition.
Augmentation. Increase beyond the
normal number; 179, 200.
Auranti-.uits. Orange-colored.
Auratus, Aureus. Golden-colored, or
yellow with golden lustre.
Jlaricle (Auricula). An ear or ear-
shaped appendage.
Auticulde (-atus). Furnished with an
auricle; 96.
Autocarpous A fruit consisting of peri-
carp alone, having no adnate parts.
Autoyo.my. Close-fertilization, the fe-
cundation of a flower bv its own pol-
len; 215, 216.
Avenius. Veinless.
Awl-shaped. Narrow, terete or some-
what so, and attenuate from a broader
base to a slender or rigid point.
Awn. A bristle-shaped appendage, such
as the beard of Rye and Barley.
Awned. Furnished with an awn.
Axil (Axilla). The angle formed on the
upper side of the attachment of a leaf
with the stem, or the point just above
this attachment ; 6.
Axillary (-dris). In or relating to an
axil; 7.
Axile, Axial (Axilis). Relating or be-
longing to the axis.
Axis. The stem; the central part or
longitudinal support on which organs
or parts are arranged ; the central line
of any body.
Bacca. A berry ; 299.
Baccate (-ntus). Berry-like ; pulpy
throughout.
ice turn. An aggregation of berries
in one flower ; 300.
Bidius. Chestnut-brown.
Bidausta. Name applied to the fruit
of the Pomegranate, with firm rind,
crowned with the lobes of an adnate
calyx, baccate within, and many-
seeded.
Banner. The vexillum, standard, or
upper petal of a papilionaceous co-
rolla; 184.
Barb. A bristle or stout hair, which is
hooked or double-hooked, or retrorsely
appendaged at the tip.
Burba. Beard.
Barbate (-atus). Bearded ; beset with
long and weak hairs.
Barbellate (-atus). Beset with shorter
and stiffer hairs or barbdlce.
Barbellulate (-atus). Diminutive of the
preceding.
Bark. The rind or cortical portion of a
stem, especially of an exogen; 76.
Basal (Basilaris). Relating to the base.
Basal-nerved. With nerves all from the
base of the leaf; 92.
Base (Basis). The extremity by which
an organ is attached to its support.
Basidia. Cells of the fructification of
Mushrooms which bear the spores.
Basifixed (-us). Attached by the base
or lower end ; 253.
Basi(jynium. Synonym of Carpophore
or Thecaphore.
Basinerved (-ius). When the ribs pro-
ceed from the base of a leaf.
Basipetal. Developing from apex to-
ward the base.
Bast, or Bass. Inner fibrous bark ; 77.
Bast-cells. The es-ential components of
bast ; long and flexible butthick-walled
attenuated cells ; 77.
Beak. A narrowed or prolonged tip.
Beaked. Ending in a beak.
Bell-shaped. Same as Campanulate ; 249.
Berried. Baccate.
Berry. A fruit, the whole pericarp of
which is fleshy or pulpy; 299.
Bi- or Bis. As a prefix to Latin words
(Greek words have Di-), two, twice,
or doubly.
Biacuntinate (-atus). Two-pointed, as
malpighiaceous hairs, fixed by the
middle and tapering to each end.
Biarticulate (-atus). Two jointed.
Biauriculate (-atus). Two-auricled.
Bibracteate <-atus). With two bracts.
Bibracteolate (-atus). With two bract-
lets.
Bicallose (-osus). With two callosities.
Bicdiinate (-atus). Two-keeled.
Biceps. | With two supports or stalks,
Bicipital. } or two-headed.
Bicolor. Two colored.
Biconjugate (-atus). Twice paired.
Bicor-nis. Two-horned.
Bicornute. Same as preceding.
GLOSSARY.
399
Bicruris. Two-legged, or with two sup-
ports.
Bidentate (-atus). Having two teeth.
(Not doubly dentate.)
Biduus. Lasting two days only.
Biennial (Biennis). Of two years' dura-
tion ; 31.
Bifarious (-ius). Two-ranked; in two
vertical rows.
Biferus. Double-bearing; fruiting twice
a year.
Bifid (-idus). Two-cleft, to the middle
or thereabout.
Biflorous (-us). Two-flowered.
Bifoliate. Two-leaved.
Bifolioiite. Of two leaflets.
Bifoi-ate (-atus). Having two open-
ings.
Biformis. Two-formed ; in two shapes.
Bifrons. With two faces or aspects.
Bifurcate (-aim). Two-forked ; t. e. of
two prongs or forks. But it may
mean bis furcatus; i.e., forked and
again forked.
Biyeminate (-atus). Twice twin ; same
as Biconjugate.
Biyener. The offspring of a cross be-
tween two generic-ally different plants.
Bijuyate (Bijuyus). Two-paired, as a
pinnate leaf of two juga or pairs of
leaflets.
Bilabiate (-atus) Two-lipped; 247.
Bildmellate (-atus), or Bilamellar. Of
two plates or lamella.
Bilobed (Bilobus), or Bilobate. Of two
lobes, or cleft into two segments.
Bilocellate. Divided into two locelli;
263.
Bilocular (-aris). Two-celled.
Bimestris. Lasting two months.
Bimus. Lasting two vears ; two years
old.
Binary (-arias). Consisting of two
members; 176.
Binate (-atus). In pairs or twos.
Bini. Twin, or two together.
B! nodal ( Binodis). Having two nodes.
Binomial Nomenclature, 346.
Bioloyy. The natural history of plants
and animals, i. e. of living things ; 1.
Bipalmate (-atus). Twice palmately com-
pound.
Biparous. Bearing two; as a cyme of
two rays or axes; 152, 155.
Bipdrtible (-ibilis). Capable of division
into two similar parts.
Bipartite (-itus). Divided almost into
two pieces ; two-parted.
Bipts. Same as Bicruris.
Bipinnate (-atus). Doubly or twice
pinnate; 103.
Bipinndtifid (-idus). Twice or doubly
pinnatitid; 100.
B'pinndtisect (-us). Twice pinnately di-
vided.
Biplicate (-atus). Twice folded or plaited.
Biporose (-osus). Opening by two pores.
Biradiate (-atus). Of two rays.
Birimose (-osus). Opening by two slits.
Bisected (-us). Completely divided into
two parts ; 99.
Biseptate (-atus). With two partitions.
Biserial (-ialis), or Biseriate (-iatus). In
two series, one above the other.
Btserrate (-atus). When serratures are
again serrate ; doubly serrate.
Bisexual. Having both stamens and
pistil; hermaphrodite; 191.
Bisulcate (-atus, Bisulcus). Two-grooved;
having two furrows.
Biternate (-atus). Twice ternate.
Bladdery. Thin and inflated.
Blade. The lamina, limb, or expanded
portion of a leaf, &c.; 85, 245.
Blastema. The budding or sprouting
part or point. First used for the axis
of an embryo; now used for the ini-
tial growth out of which any organ
or part of an organ is developed.
Bloom. Besides its use as equivalent to
blossom, it denotes the white powdery
and glaucous covering of the surface
of many fruits and leaves, of a waxy
nature.
Boat-shaped. Of the shape of a boat, of
the deeper sort, with or without a keel.
Bostrychoidal. Having the form or char-
acter of a ringlet, or Bostryx ; 157.
Bostryx. An uniparous helicoid cyme ;
156.
Both renchyma. Tissue of plants com-
posed of dotted or pitted ducts.
Botry-cymose. Racemes or any hot ryose
clusters cymosely aggregated ; 159.
Botryose (-osus), Botryoidal. Of the ra-
cemose type; 144, 145, 146, 153.
Botrys. The equivalent of Raceme ; 146.
Botuliformis. Sausage-shaped.
Brdchiate (-idtus). With spreading
arms, as branches (especially opposite
and decussate) widely diverging
Brachys. Greek for short, and used in
compounds; as, Brachypodus, short'
stalked.
Bract, Bractea. The leaves (more of
less modified) of a flower-cluster; 118,
141.
Bracteate (-eatus). Having bracts.
400
GLOSSARY.
Bracteola, Bracteole. See Bractlet.
Bracteolate(-atus). Having bractlets.
Bractlet. A bract of the ultimate grade,
as one inserted on a pedicel or ultimate
flower-stalk, instead of subtending it;
141, 142, 160.
Bracteose (-osus). Full of, or with con-
spicuous bracts.
Branches. Secondary axes, or divi-
sions of an axis; 47.
Branchlets. Ultimate branches or divi-
sions of an axis ; 47.
Breathing-pores. See Stomata, 89.
Bristle. A stiff hair, or any slender
body or outgrowth which may be
likened to a hog's bristle.
Bristly. Beset with bristles.
Biunneus. Deep brown.
Brush-shaped. See Aspergilliform.
Bryology. The botany of Mosses.
Bud. The undeveloped state of a stem
or branch, with or without leaves;
6, 40.
Bud-scales. The teguments of a bud; 40.
Bulb (Bulbus). A leaf-bud (commonly
subterranean) with fleshy scales or
cats ; 43, 62.
Biiibiceps. A stem with bulbous base.
Bulbiftrovs (-us). Bulb-bearing.
Bulbillus, Bulbulus. Diminutive bulb.
Same as
fiulblet. A small bulb, especially such as
is produced in the air, in the axil of or-
dinary leaves, or upon them ; 63.
Bulbodium. A synonym of Conn, the
"solid bulb."
Bulbo-tuber. Synonvm of Corm.
Bulbous, Bidbosus. Having bulbs or the
structure of a bulb.
Bu Unle (-cttus). Said of a puckered sur-
face (as if blistered), thrown into por-
tions which are convex and projecting
on one side and concave on the other.
Also used in specific names, in its
more literal sense for inflated.
Bursicula. A small pouch (bursa}\
such as that which encloses the disk
or gland of the caudicle of the pollin-
ium of an Orchis.
Bin siculntus. Furnished with a bursi-
cula or pouch.
Byswtceous (-eus). Composed of fine
threads, like byfsus or fine flax.
Caducous {-us}. Dropping off very early,
as the calyx of a Poppy at the time
of expansion ; 243.
Cceruleus. Sky blue, or pure blue.
iuK. Lavender color ; pale green with
whitish or gray.
Calathidium, Cdlathis. Literally a bas-
ket; a name for the head of flowers (or
better for the involucre only) of Com-
positse
Calathiform (-ormis). Cup-shaped ; of
somewhat hemispherical outline.
Cnlcar. A spur; mostly used for the nec-
tariferous one of a calyx or corolla.
Cdlcarate (-dtus). Furnished or pro-
duced into a spur.
Cdlceolate (-atus), or Calceiformis.
Shaped like a slipper or shoe
Callose (-osus). Bearing callosities
(calli), or hard protuberances.
Calvus. Bald, as an akene without
pappus.
Calycdnthemy. Name of the monstros-
ity in which the calyx imitates an
exterior corolla ; 174.
Calyciflorous (Calyciflorce), 340.
Calycine ( Calycinus). Relating to calyx.
Calyculate (-atus). Bearing bracts next
the calyx which imitate an external
or accessory calyx.
Calyculus. An involucre or involucel
imitating an additional calyx.
Cnlyptra. The hood or veil of the
spore-case of a Moss; or some cover-
ing body like it.
Calyptrate (-atus). Furnished with a
calyptra, or something like it.
Calyptriform (-ormis). Calyptra-shaped ;
as the calyx of Eschscholtzia.
Calyx. The flower-cup, the exterior
perianth ; 164
Cdnara and its diminutive Cam^rula
(chamber) are sometimes used for the
cells of a fruit.
Cambium. Old name of the viscid mat-
ter between bark and wood in com-
mon trees or shrubs in spring; now
used for the nascent structure there
forming, or Cambium layer ; 78.
Campdnulate (-atus). Bell-shaped ; elon-
gated cup-shaped or shorter, and broad
from the baceous (-us). Of hard and brittle
texture. ,
Cryptos. Greek for concealed ; whence
Ci <,ptoyamia. Cryptogamous or Cryp-
togamic plants ; 3, 335, 344.
Crypto f/amous. Pertaining to the above.
Cucullate (-atus), Cuculldris, Cuculli-
formis. Hooded, or hood-shaped,
cowled.
Culm ( Culmus). The peculiar stem or
straw of Grain-plants and Grasses; 50.
Cultrate (-atus), Cultriformis. Shaped
like a broad knife-blade.
Cuneate (Cuneatus), Cuneiform (-ormis).
Wedge-shaped ; triangular with an
acute angle downward ; 95.
Cup-shaped. In the form of a drinking-
cup.
Cupule (Cupula). The acorn-cup and
the like; 296.
Cupularis, Cupulatus. Furnished with
or subtended by a cupule or any re-
sembling body.
Cupuliferous (-us). Cupule-bearing.
Curvinerved (-ius). When the ribs of a
leaf are curved in their course ; 92.
Curviserial. In curved or oblique ranks ;
124.
Cushion. The enlargement at or be-
"neath the insertion of many leaves.
Cuspidate (-atus). Tipped with a Cusp,
or sharp and rigid point; 97.
Cut. Same as incised, or in a general
sense as cleft.
Cuticle ( Cuticula). The outermost skin
or pellicle.
Cutting. A severed portion of a plant
used for bud-propagation ; 43.
Cydneus. A clear bright blue.
Cydthiform (-ormis). Cup-shaped; in the
form of a
Cydthus. A drinking-cup, such as a
goblet or wine-glass.
Cycle. A circle. Sometimes used for
one turn of a helix or spire; 122.
Cyclical. Relating to a cycle ; or coiled
into a circle; 119, 120.
Cylindraceous. Somewhat or nearly
cylindrical.
Cylindrical (-us). Elongated and with
circular cross-section; in the form of a
cylinder.
Cymb&form or Cymbiform (-ormis).
Boat-shaped.
Cyme ( Cyma). A flower-cluster of the
determinate or centrifugal type.
406
GLOSSAKY.
especially a broad and flattish one;
151.
Cymo-botryose. When cymes are ar-
ranged in botryose manner; 159.
Cymose (-osus). Bearing cymes, or re-
lating to a cyme ; l8l.
Cymule ( Cymula). Diminutive cyme,
or a portion of a cyme; 151.
Cynarrhodium. Name of such a fruit as
that of the Rose; fleshy, hollow, and
enclosing achenia.
Cypsela. Name of an ache, ium in-
vested by an adnate calyx, as the fruit
of Composite; 295.
Cystdiih. One of the mineral and
usually partly crystalline concretions
of the cells of the epidermis of or
subjacent tissue of the leaf in various
plants, especially in Urticaceae.
Cytoblast. An obsolete name for the
nucleus of a cell of cellular tissue.
Ddctylose (-osus). Fingered, or finger-
shaped.
Dusyphyllous (-us). Woolly-leaved.
Dealbate (-atus). Whitened over (as if
whitewashed) with a white powder or
minute pubescence.
Deca. Greek for ten, compounded with
various words, such as
Decnyynia. One of the Linnaean artificial
orders; 337.
Decdgynous (-us). With ten styles or
carpels.
Decdmerous (-us). Of ten members ; 176.
Decandria. A Linnaean class with ten
stamens; 334.
Decandrous (-us). With ten stamens;
249.
Decapetalous (-us), Decasepalous, &c.
With ten petals or sepals, &c.
Deciduous (-us). Falling, or subject to
fall in season, as petals after anthesis,
and leaves (except of evergreens) in
autumn ; 243.
Diclinate (-atus), or Declined Bent or
curved downward or forward.
Decompound. Several times compound-
ed or divided ; 102, 104.
Decompositus. Decompound.
Decumbent (-ens). Reclining, but with
summit ascending; 53.
Decurrent (-ens), Decursice. Running
down into ; as where leaves are seem-
ingly prolonged below their insertion,
and so run down the stem.
Decussate (-atus). In pairs alternately
crossing at right angles.
Deduplication, Fr. Dedoublement. Same
as Chorisis; 202.
Dejinite (-itus). Of a fixed number, not
exceeding twenty ; or of a fixed or. er.
Dejinite Jnfloresctnce. Where axes of
inflorescence end in a flower; 144, 151.
Dejlexed (-us). Bent or turned abruptly
downward.
Dtjlorate (-atus). Past the flowering
state.
Defoliate (-atus). Having cast its leaves.
Defoliation, 87.
Dehiscence (-entia). The mode of open-
ing of a capsule or anther by valves,
slits, or regular lines ; 288.
Dehiscent (-ens). Opening by regular
dehiscence; 292.
Deliquescent (-ens). Dissolving or melt-
ing away, as a stem divided into
branches ; 48.
Deltoid (-oides). Having the shape < f
the Greek letter A.
Demersed (-us). Underwater; same as
submersed.
Dendritic (-icus), Dendroid (-oideus).
Tree-like.
Dendron. Greek for tree.
Deni. Ten together.
Dens. A tooth.
Dentate (-atus). Toothed ; specially with
salient teeth not turned forward ; 98.
Denticulate (-atus). Minutely toothed;
having denticulations, or diminutive
teeth.
Dem'tdate (-atus). Made naked ; stripped.
Deormm. Downwards.
Depauperate (-atus). Impoverished; as
if starved; or diminutive for want of
favorable surroundings.
Depressed (-us). Having the appear-
ance or shape as if flattened from
above.
Derma. Greek for skin or surface of. a
plant or organ.
Descending (-ens). Tending or turning
gradually downward.
Descending Axis. Primary root; 11.
Determinate. Limited in number or ex-
tent; as are the axes of determinate
inflorescence; 144, 151.
Desinens. Terminating in.
Desmos. Greek for things bound, or as if
chained together.
Dextrorse (Dextrorsus: adv. Dextror-
sum). Toward the right hand, or re-
lating to it; 51, 140.
Di, Dis. In Greek compounds, two, or
double.
Diacheniuin. Sj'nonym of Cremocarp.
GLOSSARY.
407
Diadelphia. A Linnaean (335) class
having the stamens.
Diadtljjhous (-ua) Combined by their
filaments into two sets; 250.
Diagnosis. A brief distinguishing char-
acter.
Diaiypetalce, 341.
Dudypetalous (-us). Same as polypeta-
lous, i. e. of separate petals; 244.
Dialyphyllous (-us). Bearing separate
leaves.
Diandria. A Linnaean class with per-
fect flowers having only two stamens;
334.
Diandrous (Diander, &c.). Having two
stamens; 249.
Diaphanous (-us). Letting the light
shine through.
Dicarpdlary. Composed of two carpels
or pistil-leaves ; 261.
Dichasium. A two-parted or two-rayed
cyme; 152, 155.
Dichlamydeous (-eus). Having a double
perianth ; 191.
Dichotomuus (-us). Forked in pairs ;
two-Corked.
Dichogamous (-us), Dichogamy. Her-
maphrodite with one sex earlier de-
veloped than the other in the blossom;
219.
Diclesium. Name of a fruit consisting
of an achenium within a separate and
free covering made of perianth, as that
of Mir.tbilis.
Diclinous (Diclinis). When flowers are
of separate -sexes ; 191.
Dicoccous (-us). Fruits of two cocci.
Dicotyledons, Dicotyledones. Plants of
the class marked by having two coty-
ledons ; 27, 339, 340, 344.
Dicotyledonous (-eus). Having a pair of
cotyledons; 10, 314.
Didymous (-us). Twin, found in pairs.
Didyndmia. The Linnaean class marked
bv didynamy (335), i. e.
Didynamous (-us). When a 4-androus
flower has the stamens in two pairs,
and one pair shorter than the other ;
250.
Diercsilis. Mirbel's name for a dry
fruit composed of several cells or car-
pels connate around a central axis,
and separating at maturity, as that of
Mallow.
Difformis. Of unusual formation.
Diffuse, (-usus). Widely or loosely
spreading.
Digamous (-us). Of two sexes in the
same cluster.
Digitate (-atus). Fingered ; a compound
leaf in which all the leaflets are borne
on the apex of the petiole; 101.
Diijitattly. In a digitate mode; same
as Palmately.
Digitate- Pinnate, 104.
Digynia. A Linnaean order character-
ized by having the gyncecium
Digynous. With two separate styles or
carpels; 261.
Dimerous (-us) Of two members in each
circle; 176.
Dimidiate (-atus). Halved, or as if one-
half was wanting.
Dimorphous (-MS), Dimorphic, Dimor-
phism. Occurring under two forms;
225, 234.
DicBcia. Linnaean class (355) of plants
with the flowers
Dioecious (Diozcius, Dioicous). Unisex-
ual, and the two sexes borne by dis-
tinct individuals; 191.
Di(cio-polygamous. When some indi-
viduals bear unisexual and others bi-
sexual flowers.
Dipttalous (-us). Two-petaled; 244.
Diphyllous (-us). Two-leaved ; 243.
Diplo. See Duplo.
Diplostemonous, Dijjlostemony . Having
twice as many stamens as petals or
sepals ; 177, 198.
Diplotegium. A capsule or other dry
fruit, invested with adnate calyx ; an
inferior capsule.
Dipterous -(us). Two-winged.
Diremption (-io). Syn. of Chorisis; 202.
Disciferous (-us). Disk-bearing.
Disciform (-ormis). Depressed and cir-
cular, like a disk or quoit.
Discoidal or Discoid (Discoideus). Ap-
pertaining to a disk. A discoid head
is one destitute of ray-flowers.
Disc or Disk (Discus). A word used in
several senses. The disk or disc of a
flower is a development of the torus
within the calyx, or wilhin the corolla
and stamens; 213. In acapitulum or
head of flowers it is the central part
of the cluster, or the whole of it as
opposed to a border or ray. It is the
face or surface of any organ, such as
a leaf-blade, as opposed to the mar-
gin. In vegetable anatomy, certain
round spots or markings on cell-walls
are termed discs.
Discolor. When the two faces of a leaf,
&c., are unlike in color.
Discrete (-etus). Separate ; not coales-
cent.
408
GLOSSARY.
Dlsepalous (-us) Two-sepaled ; 244.
Disk-flowers. Those belonging to the
disk, or body, and not to the margin
or rav of a eapitulum.
Dissected (-us). Deeply cut or divided
into numerous segments.
Dissepiment (-cntum). A partition in an
ovary or pericarp; 264.
Dissilient (-ens). Bursting asunder or
in pieces.
Distichous (-us). Disposed in two ver-
tical ranks ; 122.
Distinct (Distinctus). Separate from;
not united.
Distract He (-His). Carried widely apart.
Dihecous (-us). Of two thecae, or cells,-
as are most anthers ; 254.
Diurnal. Daily ; occurring in the day;
sometimes used for ephemeral.
Divaricate (-atus). Extremely divergent.
Divergent, Diveryiny (-ens). Inclining
away from each other.
Divided (Divisus). Where lobing of
segmentation extends to the base ; 98.
Dodtca Greek for twelve. Used in
DodeccKjynia. Linnaean order with flow-
ers. .,
Dodecdyynous. Having twelve styles or
distinct carpels.
Dodecdmerous (-us). Of twelve parts in
the circle.
Dodtcandria. A Linnaean class (334)
with the flowers.
Dodecdndrous. Having twelve (or from
12 to 19) stamens; 249.
Dodrantdlis. A span (about nine inches)
long.
Dolabriform (-ormis). Axe-shaped or
hatchet-shaped.
Dorsal (-alts). Relating to the dorsum
or back.
Dorsal Suture. That which answers to
the midrib of a carpel ; 261.
Dorsiferous Borne on the back.
Double Has a technical use when a
flower is said to be "double;" this
denoting one in which the leaves of
the flower are monstrously increased
mostly at the expense of the essential
organs.
Downy. Pubescent with fine nnd soft
hairs. Loosely synonymous with soft-
pubescent, tomentose, &c.
Drepdnium. A sickle-shaped cyme; 156.
Drupaceous (-eus). Resembling or relat-
ing to a drupe.
Drupe (Drupa). A stone fruit; 297.
Drupelet, Drupel (Drupeola). A dimin-
utive drupe; 297.
Drupetum. An aggregation of drupes;
300.
Duct. In vegetable anatomy, an elon-
gated cell or tubular vessel, found espe-
cially in the woody (tibro-vascular)
parts of plants.
Diimetose (-osus). Pertaining to Dume-
tum, a thicket, or Duinus, a bush.
Dumose (-asua). Bushy, or relating to
bushes.
Duplo. Twice as many. In Greek com-
pounds, Diplo.
Duramen. The heart-wood of an exo-
genous stem ; 80.
Dwarf. Of small size or height com-
pared with its relatives.
Dijclesium. See Diclesium.
E- or Ex- As a prefix to Latin words,
carries a privative meaning, as Ec<,s-
tate, without ribs, ExaWuminous, with-
out albumen.
Eared. Same as Auriculate.
Ebracteate, Ebracteolate (-erveless; no ribs
or veins visible.
Ennea. In Greek compounds, nine ;
as in
Ennen (jyni't . A Linnrcan ordinal name,
and
Enned([yn> ytl/rostomum. Name given by Des-
vaux for such an aggregate fruit as
a raspberry; 300.
Estivation. See ^Estivation.
Etcerio. Name of aggregate fruits,
especially of fleshy ones, such as a
blackberry.
Etiolated. Blanched by darkness.
Eu. Prefixed to words of Greek deri-
vation denotes very, true, or much so
Frequently used in names of sections
or other groups ; 357.
Euphylla. Foliage-leaves, or true leaves.
Eutropic (-icus). Name suggested for
twining "with the sun;" 51.
Evalmlar (Evalvis). Not opening by
valves.
Evergreen. Bearing green foliage all
the year round.
Evittatus. Not vittate.
Ex. A prefix in place of E privative
when the following part of the com-
pound begins with a vowel ; as
Exalbuminous (-osus). Destitute of albu-
men ; 14, 309.
Exdlate (-atm). Destitute of wing.
Exanthemata. Eruptive excrescenses on
the surface of leaves, &c. ; blotches.
Exaristate (-atus). Destitute of an
arista or awn.
Exasperate (-atus). Rough with hard
projecting points.
Excentric (-icus). Out of the centre;
one-^ided.
Excuirent (-ens). Running through to
the very summit or beyond ; 48.
Exif/uus. Small or mean.
Exilis. Lank or meagre.
Eximins. Distinguished, as for size or
beauty.
Exo-. In Greek compounds, external or
/ outward ; as
Exocarp ( Exocarpium). The outer layer
of a pericarp ; 288.
Exogenous. Outside growing, as the
wood of Dicotyledons; 69, 73.
Exoyens, Exogence. Exogenous Plants;
69, 340.
Exorliizce. Name equivalent to Exogen,
from
Ex< hizal (-us). The radicle not sheathed,
so the primary root in germination
/ has no covering to break through.
Exostvme (Exostoma). The foramen of
the outer coat of the ovule ; 277.
Exothecium. The outer wall of an
r anther.
Explana'e ( atus). Spread out flat.
Exsert, Exserted (Exsertus). Protruding
beyond or out of, as stamens beyond
the corolla.
Exstipulate (-atus). Destitute of stipules.
Exterior. External in the sense of outer.
But also in the flower sometimes used
in the sense of anterior.
Kxtine. Outer coat of a pollen-grain.
Extra-axillary. Beyond or out of the axil.
Extrvrse ( t.xtrorsus, Extrorsum ). Di-
rected outward: 253.
Eye. A gardener's name for an unde-
veloped bud.
Fades. Face ; the general aspect.
Falcate (-atus), and Falciform (-ormis).
Scythe-shaped or sickle-shaped ; plain
and curved, with the edges parallel.
Family In botany, synonymous with
Order; 325.
Fan-shaped. See Flabelliform.
Farina Starch.
Farinaceous (-eus). Of the nature of
starch, or containing it.
Farinose (-osus). Covered with a meal-
like powder.
Fasciate (-atus). Said of monstrous ex-
pansions of stems, giving the appear-
ance as of several stems coalescent in
one plane.
Fascicle (-icula). A close cluster or
bundle, whether of flowers, stalks,
roots, or leaves; 147, 153.
Fascicled (Fasciculatus, Fascicularis).
In a fascicle; 131.
Fastiyiate (-atus). Said of branches
when parallel, clustered, and erect.
Faux, pi. fauces. The gorge or throat
of a gamophyllous calyx or corolla;
either at the orifice, or a portion
between the limb and the proper
tube; 246.
Fdveolate (-atus), Favose (-osus). Honey-
combed ; same as Alveolate.
Feather-veined. Having veins all pro-
ceeding from the sides of a midrib.
Feathery. See Plumose.
GLOSSARY.
411
Fcmla or Fcecula. Starch-like matter.
Applied to a pistillate flower, or to a
plant producing only such flowers.
Fenestrate (-atus), Fenestralis. Pierced
with large holes, like windows.
Ferruyineous or Ferruginous (Ferru-
(jineus). Colored to imitate iron-rust.
Fertile (-His). Fruitful, fruiting, or ca-
pable of producing fruit ; as a fertile
flower is one provided with a well-
formed pistil ; 191. In English descrip-
tions, Flower fertile usually means a
pistillate or female flower. Stamens
or anthers are also said to be fertile
when polliniferous and capable of fer-
tilizing.
Fertilization. Synonym of fecundation,
as of the ovule by pollen ; 215.
Fibre (Fibra). Any fine filament; the
elementary components of wood, &c. ;
delicate roots, &c.
Fibril (-ilia) A diminutive fibre.
Fibrillate (-atus), Fibrillose (-osus).
Furnished or abounding with fibres
or fibrils.
Fibrous, Fibrose (-osus). Composed or
of the nature of fibres.
Fibro-va scalar. Consisting of woody
fibres and ducts.
Fiddle-shaped. Obovate and with a
sinus or contraction on each side.
Fidust. A Latin termination for cleft or
lobed.
Filament (-entum). The stalk or sup-
port of an anther; 165, 251. Also any
fibre-shaped or thread-like body.
Filamentous, Filamentose (-osus). Com-
posed of threads or filaments.
Filicoloyy. The botany of Ferns: re-
placed by Pteridology.
Filiform (-ormis) . Thread- shaped; long,
slender, and terete.
Filipendulous (-us). Hanging from a
thread.
Fimbria. A fringe, or dissected border.
Fimbriate (-ntus). Fringed ; bordered
by slender processes or marginal ap-
pendages.
Fimbnllate (-atus), Fimbrilliferous (-us).
Bearing FimbriUce or diminutive fringe.
Fingered. See Digitate.
Fissiparous. Multiplying by the divi-
sion of one body into two, and so on.
Fissus. Split or cleft. See Fidus.
Fistular, Fistulose (-osus). Hollow
through the whole length, as the leaf
and stem of an Onion.
Fldbellate (-atus), Flabelliform (-ormis).
Fan-shaped; much dilated from a
wedge-shaped base, and the broader
end rounded.
Flabellinerved. With radiating straight
nerves; 92.
Fldytllate (-attis), Flayellaris. Produc-
ing filiform runners (Flayella), or
runner-like branches.
Flagelliform (-ormis). Runner-like ;
long, slender, and supple like a whip-
lash or Flayellum ; 53.
Flammeus. Flame-colored.
Flavescent (-ens). Yellowish or pale
yellow.
Fl:ivus. Pale yellow or ochre-yellow.
Fleshy. Succulent; of the consistence
of flesh.
Flexuous, Flexuose (-osus). Zigzag;
bent alternately in opposite direc-
tions.
Floating. Borne on the surface of water.
Floccose (-osus). Bearing or clothed with
locks of soft hairs or wool (flood).
Flocculent. Diminutive of floccose.
Flora (Goddess of flowers). The aggre-
gate of the plants of a country or dis-
trict; or the name of a work which
systematically describes them; 369.
Floral. Belonging to the flower.
Floral Envelopes. Flower-leaves; 164.
Floret. A small flower, one of a cluster.
Floribundus. Abundantly floriferous.
Floriferous(Floi'ifer, Floriferus). Bear-
ing flowers
Florula. A small Flora; the Flora of
a restricted district.
Flos. Latin for flower. Flosplenus. A
" double " flower; that is, one in which
petals are increased abnormally, com-
monly at the expense of the androe-
, cium or the gynoecium also; 171.
Flosculus. Latin for floret.
Flower. The whole reproductive appa-
ratus in a phaenogamous plant; 16-3.
Flower-bud. An unexpanded blossom
or undeveloped cluster; 40.
Flowering Plants, 3, 344.
Flowerle'ss Plants, 3, 344.
Fluitans. Floating.
Flumatile, Fluridtilis. Belonging to a
river or running water.
Fly-traps, 113
Fcemineus. Feminine or female flower,
plant, &c. ; 191.
Foliaceovs (-eus). Leaf-like in texture
or appearance; or bearing leaves.
Foliar (Fo/iaris). Relating to leaves.
Foliation (Foliatio). Leafing out.
Foliate (-atus). Having leaves. With
Latin numerical prefix, bifoliate, tri-
412
GLOSSARY.
foliate, and so on, according to the
number.
Foliiform (-ormis). Leaf-shaped.
Foliolate (-atus). Having leaflets : their
number may be indicated by Latin
numerals, as bifoliolate, trifoliolate,
&c.; 102.
Folidum. A leaflet; 102.
Foliuse. Bearing numerous leaves.
Folium. Latin for leaf; 85.
Follicetum. A whorl or aggregation of
follicles; 300.
Follicle (-iculus). Fruit of a single carpel
dehiscent by one (the ventral) suture;
292. Name of the earlier botanists
lor any kind of capsular fruit.
Follicular (-aris). Pertaining to or like
a follicle.
Foot-stalk. Petiole, 85 ; -or Peduncle, 143.
Foramen. An aperture of any kind;
specially that of the coat of the ovule;
277.
Foraminulose (-osus). Pierced with many
small holes.
Forcipate (-atus). Like forceps or pin-
cers.
Forked. Divided equally into branches.
Fornicate (-fitus). Arched over, as by
scales (Fornices) covering the throat
of the corolla of Hound's-tongue, &c.
Foveate (-/?), and diminutive Foveo-
late. Pitted; impressed with shallow
depressions or pits, Fovece.
Fovillai. Minute granules in a liquid,
in the protoplasm of the pollen-grain,
&c.; 258.
Free. Not adnate to other organs.
Some'imes used in the sense of dis-
tinct, i. e. unconnected with others of
the same sort.
Fringed. See Fimbriate.
Frond (Frons). An old name for leaf:
employed mainly for the leaf of Ferns
and other Cryptogamia, and certain
Phaenogamous plants which serve for
fructification as well as foliage; also
for the peculiar foliage of Palms; 67.
Frondescence (-enti(t). The act of leaf-
ing. Has also been employed to
express the metamorphosis of floral
organs into foliage-leaves; 174.
Frondose (-osus). Sometimes used in
the sense of leafy; also frond-like, or
bearing fronds.
Fructification. The act or the organs
of fruiting or reproduction through
flower and seed, or their analogues.
Fructus. Latin for Fruit.
Fruit. The immediate product of fruc-
tification ; in phaenogamous plants,
the seed-vessel and contents, along
with all intimately connected acce.-.-
sory parts; 285.
Fruit-dots in Ferns. See Sorus.
Frumentaceous. Relating to grain ( Fru-
mentum).
Frustulose (-osus). Consisting of similar
pieces or Frustules (frustula).
Frutex. A shrub.
Frutescent (-ens). Shrubby, or becom-
ing so.
Fruticose (-osus). Shrubby, or relating
to shrubs.
Fruticulose (-osus). Relating to a di-
minutive shrubby plant.
Fruticulus. A minute or low shrubby
plant.
Fugacious. Falling or fading very early ;
lasting a very short time.
Fulvous (-us). Tawny; orange-yellow
and gray mixed.
Fulcra. Accessory organs, such as ten-
drils, stipules, spines, and the like.
Fulcrate (-atus). Propped, supported
by, or provided with accessory organs.
Fuliginous (-osus). Sooty-brown.
Fungiform (-ormis and Fung ill if ormis).
Mushroom-shaped.
Funyose (-osus) Spongy in texture;
fungus-like.
Funicule, Funiculus. The stalk of an
ovule or seed ; 276.
Funnelform, Funnel-shaped; 249. See
Infundibuliform.
Furcate (-afus). Forked; or divergently
branched.
Furcellatus. Diminutively forked.
Furfuraceous (-eus). Scurfy; covered
with bran-like scales or powder.
Furrowed. See Sulcate.
Fuscous (-us). Grayish-brown in hue.
Fusiform (-ormis). Spindle-shaped ;
terete and tapering gradually to each
end; 31.
Gdlbulus. The peculiar strobile of Cy-
press and Juniper, composed of up-
wardlv thickened or fleshy scales;
30-3.
G'tlea. A helmet; name given, from
its shape, to the upper sepal of Aconi-
tum, and the upper lip of certain
forms of bilabiate corolla; 247.
Galeote (-atus). Having a galea; hel-
met-shaped; 247.
Gamo-. In Greek compounds, denotes
union by the edges or coalescence.
GLOSSARY.
413
Gamopetalous. A corolla of coalescent
petals ; formerly Monopetalous ; 244.
Gamophyllous (-us). Composed of coa-
lescent leaves.
Gamosepalous. A calyx of coalescent
sepals; 244.
Geitonoyamy. Fecundation of a pistil
by pollen of another flower of the
same plant; 216.
Geminfite (-atus). Twin ; in pairs ; two
side by side.
Gemma. A bud, specially a leaf-bud.
Gemmation. Budding-growth; or the
disposition of buds.
Gemmule (Gemmulu). Diminutive of
gemma; minute and simple buds or
bodies analogous to buds ; also sv-
nonymof Plumule. Fora time used by
Endlicher and others for the ovule.
Genera. Plural of Genus ; 323.
General (-alls). Opposed to partial ; as
general involucre.
Generic. Relating to genus.
Genetic. Genealogical ; that which comes
by inheritance.
Geniculate (-atus). Bent abruptly, like
a knee.
Genitalia. The stamens an 1 pistils or
their analogues.
Genus. Kind or group superior to spe-
cies, and which with the species gives
the name to the plant; 323.
Geoblist (-asfus). A plumule which in
germination rises from underground,
such as that of the Pea.
Germ. A growing point or initial growth,
as of a bud; or the Embryo; 311. Or
in the sense of
Germen. The Linnaean name of the
ovary; 166.
Germination (-atio). The act of devel-
opment of the embryo of a seed into a
plant.
Gerontoyaious (-ceus). Belonging to the
Old World.
Gibbous, Gibbose (-onus) Swelling out
on one side into a (jibber or yibber-
outy,
G! (/ante us. Of unusual height.
Gilms. Dirty yellow with a tint^e of red.
Glabrous (Glnber). Smooth in the sense
of not pubescent or hairv.
Glabrate (-atus). Somewhat glabrous,
or becoming glabrous.
Glabriusculus. Almost but not quite
glabrous.
Gladiate (-atus). Sword-shaped ; in the
form of a sword-blade,whether straight
or somewhat curved. See Ensiform.
Gland for Glans. An acorn and the
like; 206.
Gland (Glandula). A definite secreting
surface or structure on the surface of
any part of a plant, or partly imbedded
in it, extended to any protuberance
or structure of similar nature which
may not secrete.
Glandular, Glandulose (-osus). Bearing
glands or having the nature of glands.
GlnnduUferous (-us). Gland-bearing.
Glarevsus. Growing in gravel.
Glaucescent (-ens). Verging upon or be-
coming glaucous.
Glaucous (-us). Covered or whitened
with a bloom, like that on a Cabbage-
leaf.
Globose (-osus). Having or approaching
a spherical form,
Globular (-arts), Globulose (-osus). Some-
what or nearly globose.
GlocJiideous, Glochidiate (-atus). When
bristles and the like are barbed at tip.
Glochis. A barb.
Glomerate (-atus). Compactly clustered,
especially into a
Glomerule ( Glomerulus, Glomus). A
cyme condensed into a head or capi-
tate cluster; 152.
Glossology, 3, 359.
Glumaceous (-eus). Pertaining to or re-
sembling glumes.
Glume, Gluma. One of the chaff-like
bracts of the inflorescence of Grasses
and their relatives ; 143.
Glumella. Diminutive of gluma ; an
inner or secondary glume.
Glutinous ( -osus). Covered with a sticky
exudation.
Gonophore ( Gonophorum). A stipe which
elevates both stamens and pistil; 212.
Gossypine (-inus). Cottony; flocculent.
Grdtilis. Slender.
Gram. See Caryopsis.
Gramineous (-em). Relating to grass or
grain-bearing plants.
Granular (-cms), Granulose (-osus).
Composed of small grains or Granules.
Granulate (-atus), Granuliferus. Bear-
ing grains or grain-like bodies.
Grareolens. Unpleasantly strong-scented.
Griaeus. Gray or bluish-gray.
Grumous (Grumosus). Consisting of
clustered grains.
Guttate (-atus). Spotted as if by drops.
Gymndnthous (-us). Naked flowered.
Gymnos. Greek for naked ; used in com-
pounds such as
Gymnocdrpous (-us). Naked-fruited.
414
GLOSSARY.
Gymnotpermia. A Linnsean artificial
order of Didynamia, in which the
nutlets resulting from tour divisions
of an ovary were taken for naked
seeds; 337.
Gyrnru.sperms, Gymnospermce. A sub-
class of naked-seeded plants ; 268, 344.
Gymnospermous (-us). Naked-seeded, as
opposed to Angiospermous.
Gynandria. A Linnaean class, character-
ized by the flower being
Gynandrous. Stamens borne on (adnate
to) the pistil, even to the style or
stigma; 251, 335.
Gynobfise ( Gynobasis). An enlargement
or production of the torus on which
the gyncccium rests or is somewhat
elevated; 212..
G yno-dioRcious. Dioecious with some
flowers hermaphrodite and others pis-
tillate only; 191.
Gijnoscium. The pistil or collective pis-
tils of a flower; the female portion of
a flower as a whole; 165.
Gynophore (Gynophorum). The stipe of
a pistil; 212.
Gynosteyium. A sheath or covering of
the gynoscium, of whatever nature.
Gynostemium. The column of an Orchid,
consisting of androscium and summit
of the gyncccium combined.
Gyrate (-atus). Curved into a circle, or
taking a circular course.
Gyrose (-osus). Curved backward and
forward in turns.
Habit (Habitus'). The general appear-
ance of a plant.
Habitat. Habitation ; the geographical
limits or station; 366.
Hcematitic (-icus). Brown-red.
Hairs. Outgrowths of the epidermis,
consisting of single elongated cells, or
of a row of cells.
Hairy. Descriptively applied to pilosity
or pubescence, in which the hairs are
separately distinguishable.
Halbert- or Halberd-shaped. See Has-
tate.
Haloed. See Dimidiate; with one half
absent or appearing to be so.
Hamate (-atus). Hooked at the tip.
Hdmulate or Hamulose (-osus). Dimin-
utive of Hamate.
Haplos. In Greek compounds, simple
or simply, as
Haplopetalus (-us). With only one row
of petals.
Haplostemonnus (-us). With a single
series of stamens; 177.
Hastate (-atus), Hastilis. Halberd-
shaped, like the head of a halberd,
i. e. sagittate, but the basal lobes di-
rected outward or at right angles to
the midrib of the leaf; 96.
Head. The form of inflorescence termed
Capitulum, viz. a cluster of sessile
flowers on a very short axis and centri-
petal in evolution; 147.
Heart-shaped. Ovate with a sinus at
base; 96.
Heart-wood. The older and matured
wood of an exogenous stem ; 80.
Hebetate (-atus). Having a dull or blunt
and soft point.
Helicoid (-oideus], Heiicoidal. Coiled into
a helix, or like a snail-shell. In true
helicoid inflorescence, the flowers are
all in a single row; 155, 157.
Helmet. See Galea.
Ifelvulus. Dull and grayish yellow.
Hemi. Half or halved; in Greek com-
pounds, such as
Hemi-anatrapous. Half anatropous.
Hemicarj) (-arpium). Half or one carpel
of a Cremocarp.
Hemitropous (-us). Same as amphitro-
pous or half anatropous.
Hepta. The Greek numeral seven.
Heptayynia. A Linnaean artificial order,
having seven styles or distinct car-
pels ; 337.
Hrptdmerous (-us). Of seven members.
Heptandria. The Linnaean class with
seven stamens ; 334.
Heptandrous. Seven-stamened; 249.
Herb (Herba). A plant with no persist-
ent woody stem above ground ; 50.
Herbaceous. Of the texture, color, or
other characters of an herb.
Herbarium, Herbal. A collection of
dried specimens of plants, systemati-
cally arranged ; 380.
Hercoyamous (-us). Said of hermaphro-
dite flowers when some structural ob-
stacle prevents autogamy.
Hermaphrodite (-itus). Of both sexes;
191.
ffetperidium. A hard-rinded berrv, like
an orange and lemon ; 299.
Heteracmy. Synonym of Dichogamy ;
219
Heteros. In Greek compounds, denotes
diverse or various, as
IletiT'icmpous (-us, Heterocarpicus).
Producing more than one kind of
fruit.
GLOSSARY.
415
Heterocephalous (-us). Bearing two
kinds oi' head or capitulum.
Heteroclite (-itus). Anomalous in forma-
tion.
Heterocline (-inus). Nearly same as
Heterocephalous ; on separate recepta-
cles.
Heterodromous (-us). Spirals of changing
direction.
Heteroyamous (-us). Bearing two kinds
of flowers.
Heterogeneous. Not uniform in kind.
Heteroyone or Heteroyoneus. When the
flowers are dimorphous or trimor-
phous as respects relative length, &c.,
of stamens and pistil ; 225, 234.
Heterosfylecl. Same as Heterogone; 234.
Heteromerous. Of members not corre-
sponding in number.
Heterophyllous (-us). Having leaves of
more than one form.
H eterotropous (-us). Turned in more
than one direction, or in an unusual di-
rection; same as Amphitropous ; 279.
(Also used by Agardh for collateral
ovules turned back to back; 282.)
Hexa. Greek numeral six ; from which
is formed
Hexayynia. Linnaean artificial order, of
flowers with six styles or distinct car-
pels ; 337.
Htxdf/ynous. Having the character of
Hexagynia.
Hexdmerous (-us). Of six members; 176.
Ilexandria. Linnaean class with perfect
flowers of six stamens ; 334.
Hexdndrous. Having six stamens; 249.
Ihxaj)etalous (-us). Having six petals.
Jlexaphyllous. Six-leaved.
Ih-xupterous (-us) Six-winged.
ffexctsrpalous. Having six sepals.
Hexafte'monous. Having six stamens.
I/ibernaculum. A winter-bud; 40.
J/idden. Concealed from sight; as
Hidden-veined, where the veins are in-
visible, as in the leaves of Pinks and
Houseleeks.
Hiemal (-nlis). Relating to winter
Hilar (Hilaris) Belonging to the hilum.
Hilum. The scar or place of attachment
of the seed ; 277, 305.
Hippocrepiform (-ormis). Horseshoe-
shaped.
Hirsute (~utus). Pubescent with rather
coarse or stiff hairs.
Hirtellous (-us). Minutely hirsute.
Hirtus. Hairy, nearly same as Hirsute.
Hispid (-idus). Beset with rigid or
bristlv hairs or with bri.-tles.
Hispidulous (-us). Minutely hispid.
Hoary. Grayish-white with a fine and
close pubescence. See Canescent.
Holose riceous (-etis). Covered with fine
and silky pubescence.
Homocnrpous (-us). With fruit all of one
kind.
Honwdromous (-us), Homodromy. With
spirals all of uniform direction.
Homoyamous (-us). Bearing one kind
of flowers.
Homogeneous. All of one nature or kind.
Hombyonous or Homoyone. Homomor-
phous as respects the stamens and pis-
til; opposed to dimorphous; 225.
Homoloyue. A homologous organ or
part.
Homologous. Of one name or type, such
as leaves and parts answering morpho-
logically to leaves; 6.
Homomdllus. Said of leaves and the like
which are all turned in one direction.
Homomorphous (-us). All of one form.
Jlomostyled. Same as Homogone.
Homotropous (-us). Curved or turned in
one direction ; applied also to the em-
bryo of an anatropous seed, with rad-
icle next the hilum ; 312.
Hood. See Cucullus.
Hooded. Bearing or in form of a hood.
Hornus, Hornotinus. Of the present
year.
Horny. Of the consistence of horn. See
Corneus.
Hortensis, Hortuldnus. Pertaining to
the garden.
Hortus siccus. Old name of an herbarium.
Humi. On the ground.
Humifusus, Humistrdtus. Spread over
the surface of the ground.
Humiiis. Low of stature.
Hyaline (-inus). Transparent or trans-
lucent.
Hybrid. A mongrel, or cross-breed of
two species; 321.
Hydrophytes (HydropJtyta). Water-
plants.
Hyemdlis. See Hiemalis.
Hypanthium. An enlargement or other
development of the torus under the
calyx : 214.
Hypo. In Greek compounds, denotes
under, beneath, lower.
II i ipn nth odium. Sa^i e as Syconium;
149, 303.
Hypoclilium. The basal portion of the
labellum of an Orchid.
Hypocraterimorphous (-us), or Hypocra-
teriform, but the latter is a hybrid of
416
GLOSSARY.
Greek and Latin. Salverform or sal-
ver-shaped; that is, in the form of a
salver raised on a central support or
stem beneath. Said of a corolla and
the like with slender tube abruptly ex-
panded into a flat limb ; 248.
Hypoyceous (-ceus). Growing or remain-
ing' underground; 19.
Hypoyynous (-us). Under or free from
the gynoecium or pistil ; 182.
Hypophyllous (-us). Growing on the un-
der side of a leaf.
Hypophyllium. An abortive leaf or scale
under another leaf, or seeming leaf, as
in Asparagus and Ruscus.
Hypsophylla. Answers to the German
" Hochblatter," or high leaves, those
of the inflorescence, L e. bracts and
the like; 6.
Hysteranthous (-us). With leaves pro-
duced later than the blossoms.
Icosandria. The Linnaean class with
twenty stamens (as the name denotes)
or a larger number, inserted on the
calyx ; 334. fcosandrous is the corre-
sponding adjective; 249.
Imberbis. Not bearded.
Imbricate (-atus), Imbricative. Over-
lapping so as to " break joints," like
tiles or shingles on a roof; either with
parts all in one horizontal row or cir-
cle, as in the aestivation of a calyx or
corolla, when at least one piece must
be wholly external and one internal;
or with the tips of lower parts covering
the bases of higher ones in a succession
of rows or spiral ranks ; 135.
Immarr/inate (-atua). Not margined or
bordered.
Immersed (-us). Growing wholly under
water.
Impari-pinnnte. Pinnate with an odd
terminal leaflet; 101.
In (equilateral (-alls). Unequal-sided.
Iri'inis. Empt}', as an anther containing
no pollen.
Inappendiculate (-atus). Not appen-
daged.
Incanescent. Same as Canescent.
Incanus Hoary-white.
Incarnate (-atus). Flesh-colored. See
Carneus.
Incised (-us). Cut irregularly and
sharply; 98.
Included (Inclusus). When the part in
question does not protrude beyond the
surrounding organ.
Incomplete (-us). Wanting some essen-
tial component part ; 190.
Incrassate (-atus). Thickened.
Incubous (-us). The tip of one leaf or
other part lying flat over the base of
the next above it.
Incumbent (-ens). Leaning or resting
upon.
Incumbent Anther. One lying against
the inner face of filament ; 253.
Incumbent Cotyledons, when the back
of ot.e lies against the radicle; 313.
Incurved (-us). Bending from without
inward.
Indefinite (-itus). Relates usually to
number, this either uncertain or too
many for easy counting.
Indefinite Growth, 49.
Indefinite Inflorescence, same as Inde-
terminate; 144.
Indehiscent (-ens). Not openingby valves,
chinks, or along regular lines; 288.
Indeterminate. Not terminated abso-
lutely, as the inflorescence in which
no blossom ends the axis of the flower-
cluster; 144, 146.
Indigenous (-us). Native and original
to the country.
Individuals, 315.
Jndivisus. Undivided, i. e. not cleft,
lobed, or parted.
Indumentum. Any hairy covering or
pubescence which forms a coating.
Induplicate (-atus). With edges folded
in or turned inward.
Indusium. The proper (often shield-
shaped) covering of the sorus or fruit-
cluster of a Fern.
Induviate (-atus). Clothed with with-
ered parts or Induvice (clothing).
Inequilateral. Unequal-sided; 106.
Inmnis. Unarmed, without prickles,
thorns, &c.
Infei~ior (InJ'erus). Said of one organ
when below another. In the blossom
also in the sense of anteiior; 160. An
inferior calyx is one below the ovary,
or free; 183. An inferior ovary is one
with adnate or superior calyx ; 183.
Inflated (-atus). Bladdery.
Inflexed (-us). Bent or turned abruptly
inward.
Inflorescence. Mode of disposition of
flowers ; less properly used for a flower-
cluster itself; 141.
Infra-axillary (-aris). Below the axil.
Infundibuliform (-ormis), Infundibular
(-aris). Funnelform, funnel-shaped;
249.
GLOSSARY.
417
Innate (-atus). Borne on the npex of
the supporting part; in an anther the
counterpart of adnate ; 252.
Innovation (-io). A new-formed shoot.
Inosculating. Same as Anastomosing.
Inseparate, Insejmration. Terms pro-
posed by Masters to express coales-
cence; 181.
Inserted (-us). Attached to or growing
out of.
Insertion (-io). Is the mode or place
where one body is attached to that
which bears it.
Integer. Entire in the sense of un-
divided, or not lobed ; 97.
Integerrimus. Entire in the sense of
quite entire, i. e. the margin without
dentation; 97.
Inter. Between; as in Intercellular,
between the cells, &c.
Interfoliaceous (-eus). Between the
leaves of a pair, as the stipules of
many Rubiaceae.
Intemodt (-odium). The portion of
stem between two nodes ; 6.
Interpetwlar (-arts). Between the pet-
ioles.
Interruptedly pinnate. Pinnate with-
out a terminal leatiet.
Inline. The inner coat of a grain of
pollen.
Intrafoliaceous (-eus). Within or be-
fore a leaf.
Introjlexed (-us). Same as inflexed.
Introrse (-o)-sus). Turned inward or
toward the axis ; 253.
Introvenius. Same as hidden-veined.
Intruse (-usus). Pushed or projecting
inward.
Inoolucellate (-atus). Provided with a
secondary involucre or
Jnvolucel (-ellum). An inner or secon-
dary involucre that of an umbellet,
&e.; 142.
Involucrate (-atus). Provided with an
involucre.
Involucre (Involucium). A circle of
bracts subtending a flower-cluster;
142.
Involute (-utus). Rolled inward; 133.
Irregular (-(tris). Exhibiting a want of
symmetry in form; 184.
Irregularity, 179, 184, 219.
Isadelphous (-us). Equal brotherhood, as
when the number of stamens in two
phalanges is equal.
Isochrous. All of one color or hue.
Isomerous (-us). The members of suc-
cessive circles equal in number; 175.
Isostemonous (-us). The stamens just as
many as the petals, &c. ; 177.
fsostemony, 196.
Jointed. See Articulated.
Juba. A loose panicle, with axis de-
liquescent.
Juyum, pl.juya. A pair of leaflets. So
pinnate leaves are unijuyate, with a
single pair of leaflets; bijuyate, with
two pairs; trijuyate, with three pairs
or juya, &e. Also the ridges on the
fruit of Umbelliferae are termed juga.
Julus. Same as Amentum or Catkin.
Julaceous (-eus). Catkin-like, Amen-
taceous.
Keel. A central dorsal ridge, like the
keel of a boat. The two anterior
petals of a papilionaceous corolla,
which are united into a body shaped
like the keel or the prow of a vessel;
185.
Keeled. Having a keel. See Carinate.
Kernel. The nucleus of an ovule, or
of a seed, i. e. the whole body within
the coats.
Kermesinus. Of the color of carmine.
Key-fruit. See Samara; 294.
Kidney-shaped. Crescentic with the
ends rounded ; very oblately cordate ;
96.
Kingdom, 325.
Labe'llum. One of the petals of an Or-
chideous flower, which is unlike the
others.
Labiate (-us). Lipped, mostly Bilabiate ;
247.
Labiatiflorous (-us). Said of certain
Composite with bilabiate corollas.
Labiose (-osus). Said of a polypetalous
corolla which has the appearance of
bilabiation.
Labium. See Lip.
Laierate (Lacerus). Irregular cleft as
if torn or lacerated.
Lacinia. A slash; used for a slender
lobe.
Laciniate (-attis). Slashed; cut into
narrow incisions.
Licinula. A diminutive lacinia or nar-
row lobe.
Lactescent (-ens). Yielding milky juice.
Lacteus. Milk-white.
Ldcunose (-osun). Abounding in pits,
holes, or depressions (lacunae).
Lacustrine (Lacustris). Belonging to Of
living in lakes or ponds.
418
GLOSSARY.
levigate (-atus). Smooth as if polished.
Lcevis (this form, and not levis, lias al-
ways been used in botany). Smooth
in the sense of not rough.
Lageniform (-ormis). Shaped like a
Florence flask or a gourd (the fruit of
Lagenaria).
Layopus. Hare-footed. Densely cov-
ered with long hairs.
Lamella. A thin plate.
Lamellar (-am), Lamellate (-atus), La-
mellose (-onus). Composed of thin
plates or lamella 1 .
Lamina. The blade or expanded part
of a leaf, &c ; 85, 245.
Lanate (-atus), Lanose (-osus). Bearing
long and implexed hairs or wool
(lana).
Lanceolate (-atus). Shaped like a lance
or spear-head; narrower than oblong,
and tapering to each end, or at least
to the apex ; 95.
Lanuyinous (-osus). Cottony or woolly ;
clothed with soft and implexed hairs
or down (lanugo).
Lappaceus. Like a bur (lappa).
Lasianthus. Woolly-flowered .
Latent. Undeveloped or dormant, as
certain buds ; 40.
Lateral (-alts). Belonging to or borne
on the sides.
Lateritious (-ius). Of a brick-red color.
Latex. Proper juice, milky juice, and
the like.
L-ttidferous. Containing or conveying
latex.
Latiseptus. With broad partition.
Lavender-color. Pale blue with some
gray.
Laxus. Loose.
Leaf. The principal sort of appendage
or lateral organ borne by the stem or
axis; 85.
Leaf-blade. The lamina of a leaf.
Leaf-bud. A bud which develops into
a leafy branch or its continuation; 40.
Leaflet. A blade or separate division
of a compound leaf ; 100.
Leaf-scar. The cieatrix left by the ar-
ticulation and fall of a leaf; 47.
Leafstalk. A petiole or footstalk to a
leaf-blade; 85, 104.
Leathery. See Coriaceous.
Lecus. A synonym for Conn.
Legume (Legumen). The seed-vessel of
Legnminosae, a carpel which normally
dehisces by both the ventral and the
dorsal suture; 292.
Pertaining to a legume,
or to the order to which the legume
gives its name.
Lenticels (Lenticellce). Lenticular spots
on young bark.
Lenticular (-ariii). Lens-shaped, that is
the shape of a lentil or a double-convex
lens.
Ltntiginosus. Covered with minute dots
or freckles.
Ltpal, Lt-palum. A made-up word to
signify a stamen transformed into a
scale, nectary, &c.
Lepicena. Unused name for a glume of
Grasses.
Lepidote (-otus) Beset with small scurfy
scales.
Lepis. Greek term for a scale.
Leptos. Greek for slender; as in Lep-
tophyllus, slender-leaved.
Leucos. Greek for white; whence
Leucanthus. White-flowered.
Leucophyllus. White-leaved, &c.
Liber. The inner and often fibrous
bark ; 77, 81.
Lid. See Operculum.
Ligneous (-eus), Liynosus. Woody.
Ligule (Liyula). A strap or strap-
shaped body, such the principal part
of a ray corolla in Composite. The
thin and scarious projection from the
summit of the sheath of the leaf of
Grasses, &c. ; 106. Or a similar out-
growth of the inner face of certain
petals; 211.
Lif/ulate (-atus), Liyuliform (-ormis).
Furnished with a ligule; 148, 247.
Liyul'iflorous (-us). Said of the head of
those Composite which contain only
ligulate corol as.
Liliaceous. Lily-like; 246.
Limb ( Limbus). A border, /. e. the ex-
panded part of a gamophyllons peri-
anth, &c., as distinguished from the
tube and throat ; 245. Sometimes the
term is applied to the lamina or blade
of a petal or a loaf.
Limbite f-itus}. Bordered.
Line (Linen}. The twelfth part of an
inch. By some reduced to the tenth
of an inch ; but the decimal line is un-
usual in botanical measurement.
Linear (-aris\. Narrow, several times
narrower than wide and the margins
parallel; 95.
Lineate (-atus). Marked with lines.
Lineolate (-atus). Marked with fine or
obscure lines.
Lingufeformis or Linguiformis, also Lin-
(julate (-atus). Tongue-shaped.
GLOSSARY.
Lip. One of the two divisions of a bila-
biate corolla or calyx, i. e. of a gamo-
phyllous organ which is cleft into an
upper (superior or posterior) and a
lower (inferior or anterior) portion or
lip (labium).
Litoral or Littoral {-alts}. Belonging
to or growing on the seashore or river-
shore.
Livid (-idus). Pale lead-colored.
Lobe, (Lobus). An}' division of an organ ;
or specially a rounded division or pro-
jection; 98.
Lobate (-atus) or Lobed. Divided into
or bearing lobes ; 98.
Lobulate (-atus). Divided into small
lobes. Lobelets, or Lobules ; 98.
Locellate (-atus). Divided into loctlli;
263.
Locellus. A secondary cell ; as where
a proper cell (loculus) of an anther or
an ovary is divided by a partition into
two cavities ; 251, 263.
Loculament (-entum). S;.me as Loculus;
289.
Locular (-arts). Celled; as bilocular,
two-celled; trilocular, three-celled,
quadrilocular, four-celled, &c.
Loculicidal (-idus), Loculicide. Dehis-
cent into the cell or cavity of a peri-
carp by the back, i. e. through a dor-
sal suture ; 289.
Loculus. The cell or cavity in an ovary
or an anther.
Loculose (-osus). Partitioned off into
cells.
Locusta. Name of a spikelet in Grasses.
Lodicule, Lodicula. One of the small
scales next to the stamens in the flower
of Grasses.
Lomentaceous (-eus). Bearing or re-
sembling a
Loment (Lomentum). A legume which
is constricted or which separates into
one-seeded articulations ; 293.
Lorate (-atus). Strap-shaped or thong-
shaped ; same as much-elongated
linear.
Lucid (-idus). With a shining surface.
Lunate (-atus). Half moon-shaped;
crescent-shaped
Lunulate (-atus). Diminutive of Lunate.
Lupuline (-inus). Resembling a head of
Hops.
Lurid (-idus). Dingy-brown.
Lusus. A "sport" or variation from
seed or bud ; 319.
Luteolus. Yellowish; diminutive of
Luteus.
Lutescent (-ens). Becoming yellow, or
faintly yellow.
Luteus. Latin for yellow.
Lycotropous (-us). Said of an ortho-
tropous ovule when bent into an open
curve or horseshoe form.
Lyrate (-atus). Lyre-shaped; a pinnat-
ilid form with terminal lobe large and
rounded and one or more of the lower
pairs small ; hence Lyrately pinnate ;
101, &c.
Macros. Greek for large or more prop-
erly long; hence Macranthus, long-
flowered ; Macrocephalus, large-
headed; Macropodus, long-footed, or
with long stalk, &c.
Mdcrospore. The larger kind of spore
in Lycopodiaceae, &c.
Maculate (-atus). Spotted or blotched,
i. e. with maculce.
Malpiyhiaceous Hairs. Those fixed by
the middle and tapering both ways.
Mdmillate (-atus), Mamillar (-aris).
Bearing teat-shaped processes.
Mammceform (-ormis). Breast-shaped
or teat-shaped ; conical with rounded
apex
Mammosus. Breast-shaped.
Mancus. Deficient or wanting.
Mdnicate (-atus). Said of pubescence
so dense and interwoven that it may
be stripped off like a sleeve.
Marcescent (-ens), Marcidus. Withering
without falling off; 243.
Maryinate (-atus). Furnished with a
margin of a distinct character or ap-
pearance.
Maryinicidal. Dehiscent by the dis-
junction of the united margins of car-
pels; 290.
Marmoratus. Marbled ; traversed by
veins or shades of color.
Maritime (Maritimus). Pertaining to
the sea or seacoast.
Mas, Masculus, Masculinus. Belonging
to the stamens, or staminate plant, or
flower; 191, &c.
Masked. See Personate.
Mealy. See Farinaceous.
Medial, Median (Medidnus) Belong-
ing to the middle ; in the plane of
bract and axis; 160.
Medifxus. Fixed by the middle; 253.
Medulla. Pith; 75."
Medullary. Relating to the pith Med-
ullary Rays, 74 ; Medullary sheath, 75.
Meiostemonous. With fewer stamens
than petals.
420
GLOSSARY.
Malleus. Having the taste or smell of
honey.
Mtttiyo. Honey-dew.
Mtnibranoui, Membranaceoug (-eus).
Thin and rather soft or pliable, like
a membrane.
Meniscoid (-oideus). Concavo-convex,
like a meniscus.
Mericarp (-arpium). One of the akene-
like carpels or a closed half-fruit of
Umbelliferaj ; 297.
Merismatlc. Dividing into parts or sim-
ilar portions.
Merithallus. Synonym of Internode.
Merom (-us). In Greek compounds, de-
notes parts or members; hence Dime-
rous, of two parts, &c.
Mesocarp (-arpium). The middle layer
of a pericarp ; 285.
Mtsophlceum. The middle or green
bark; 76.
Metamorphosis, Metamorpny, 167.
Micropyle (-yla). The spot or point in
the seed at which was the orifice of the
ovule; 277, 305.
Microspore. The smaller kind of spore
in Lycopodiacese, &c.
Midrib. The central or main rib; 92.
Miniate (-atus). Vermilion-color.
Mistus or Mixtus. A cross-breed ; 321.
Mitrceform or Milriform (-ormis). Mi-
tre-shaped or cap-shaped.
Monadefphia. The Linmean class con-
taining flowers with Monadelphous
stamens, i. e. those united by their
filaments into a tube or column ; 250,
335.
Monandria. The Linnacan class (334)
containing flowers with Monundrous,
that is, a single stamen ; 249.
Mondnthous (-us). One-flowered.
Moniliform (-oi'mis). Necklace-shaped ;
cylindrical and with contractions at
intervals.
Monocarpellary. Of one carpel ; 261.
Monucdrpic (-icus). Monocarpous, Mon-
ocarpirm. Only once fruiting; 33.
Monucephalous (-us). Bearing a single
capitulum.
Monochaslum. A cyme with one main
axis; 152, 155.
Monochlamydeous (-eus). Having but
one kind of perianth; 190, 340.
Monoclinous (-us). Synonym of Henna?
phrodjte.
Monocotyledon, Monocotyledones, adj.
Jfonocotyledonous (-eus). Plants or em-
bryo with a single cotyledon 5 23, 27,
314, 339.
Monolocular (-aris). One-celled.
Munwcia. Name of Linnsean class (335)
with flowers.
MonJiolof/y, 5.
Moschafe (-atus). Exhaling the odor of
mupk.
Mucilftyinotis, Mucildf/inosus. Slimy;
of the consistence or appearance of
mucilage.
Mucro, Mucronation. A short and
abrupt small tip to a leaf, &c.
Mucronute (-atus). Tipped with a mu-
cro; 97.
Mucronulnte (-atus). Minutely mucro-
nate.
Mule. A hybrid or cross-breed.
Miiltlcipital ( Muhiceps). Man} r -headed \
many shoots pr stems from the prowi}
of one roof.
GLOSSARY.
421
Multifarious (-us). Many-ranked, as
It-aves in several vertical ranks.
Multl/id (-idm). Cleft into many lobes
or segments.
Multiflorous (-us). Many-flowered.
Multljiiyate (Multijuyus). In many
pairs or juga.
Multilocular (-(iris). Many-celled or
several-celled.
Multiparous. Many-bearing; said of a
several-branched cyme ; lb'2, 155.
Multiple fruits. The fructification of a
flower-cluster when confluent into one
mass; 301.
Multiplication. Same as Augmentation ;
179, 200.
Multiserial (-alls), Multiseriate (-atus).
In several series.
Muricate (-atus). Rough with short and
firm excrescences.
Muriculate (-atus). Minutely muricate.
Muscariformis. Fly-brush-shaped.
Musclform (-ormis). Moss-like in ap-
pearance.
Muscoloyy. The botany of Mosses. But
is a hybrid word, and is replaced by
Bryology.
Muticous (-us). Pointless, blunt, awn-
less.
Mycelium. The filamentous vegetative
growth of a Fungus.
Mycoloyy, Mycetoloyy. The botany of
Fungi.
Xfycropyle. Micropyle misspelled.
Naked. Wanting some usual covering;
as flowers without perianth, ovules
without coats, seeds not in a pericarp,
buds without scales.
Nn pi. form (-ormis). Turnip-shaped; 31.
Nanus. Dwarf.
Nntant (-am). Floating or swimming
under water.
Nttviculir (-ana). Boat-shaped. Same
as Cymbiform.
NebuloKe (-osus). Clouded or misty.
Neck. See Collum.
Necklace-shaped. See Moniliform.
Nectar. The sweetish secretion by va-
rious parts of the blossom from which
bees make honey.
Nectary (Nectarium). The place or
thing in which nectar is secreted :
formerly applied also to any anoma-
lous part or appendage of a flower,
whether known to secrete honey or
not ; especially' to the hollow spurs of
a Violet, Larkspur. Columbine, and
the like.
Nectariferous (-us). Nectar-bearing.
Needle-shaped. See Acerose.
Nemorosus, Nenwralis. Inhabiting
groves.
Nervation. Same as Venation, or un-
branched venation.
Nerve (Nervus). In botany, this is a
simple or unbranched vein, or a slen-
der rib.
Nerved, Nervose (-osus), Nervate (-atus).
Having nerves ill the botanical
sense.
Nervulvse (-osus). Diminutive of ner-
vose.
Netted. Same as Reticulated ; Netted-
vc-ined; 92.
Neuiuse (-osus). Same as Nervose.
Neura being the Greek for nerve.
Neuter, Neutral. Sexless ; as a flower
which has neither stamen nor pistil;
]91, 195.
Nlyer. Black or blackish.
Niyricans. Turning black or verging
to black.
Nitidus. Smooth and shining.
Nicdlis. Growing in or near snow.
Niceus. Snow-white.
Nodding. Hanging down.
Node (Nodus). Literally a knot; the
portion of a stem which normally
bears a leaf or whorl of leaves ; G
Nodose (-osus). Knotty or knobby.
Nodulose (-osus). Diminutive of Nodose.
Nomenclature, 3, 345.
Normal (-ear-
ing progeny, in the way of offshoots.
Proliferation or
Prollficnt'mn is usually taken as the
production by one organ of something
different, such as the development of
buds and plantlets on leaves, of leafy
shoots in place of flowers, &c. : 73.
Prolif/erous (-us). Same as Proliferous.
Prone (Promts). Lying flat, especially
face downward.
Prvpdcultim, Propdyulum. Name of a
shoot, such as a runner or sucker which
may serve for propagation.
Propt'taines. Same as Bulblets.
Prophylln. Primary leaves, as the first
leaves of a b - anch or axis.
Prosenchyma. Plant-tissue consisting of
lengthened, tubular, or fusiform cells.
Prostrate (-atus). Lying quite flat rn
the ground; 53.
Protos. Greek for first ; used in various
compounds, such as
Protandrous, Protandry. See Proter-
androus.
Proterdndrous, also Protandrous, Pro-
terandry. When the anthers of a
flower are in anthesis earlier than the
stigma; 219, 220.
Proteranthous (-us). Where flowering
precedes leafing.
Proteroyynous, Proteroyyny, or Pro-
toyynous, Protoyyny. When the
stigma is ready for its functions ear-
lier than the anthers of the same
blossom; 219.
Protophytes, Protophyta. Alga, c.,
the supposed first plants.
Protoplasm, Protopldsma. The forma-
tive organic material of plants and
animals, in its living state.
Pruinate (-atus), Pruinose (-osits). As if
frosted over with a bloom or powder.
Pseudos. Greek for false, a prefix in
various compounds, as Pseudo-mono-
cotyledonous ; 26.
Pseudo-bulb. A thickened and bulb-
like internode in epiphytal orchids ; a
corm.
Pseudocarp (-a/pitim). The principal
or accessory part of an anthocarpous
fruit; 300."
Pseudo-costate False-ribbed, as where
a marginal or intramarginal vein or
rib is formed by the confluence of the
true veins.
Pseudospermium. Name given to any
kind of "lie-seeded fruit which is inde-
hiscent and resembles a seed, such as
an akene, &c.
Psilos. Greek for naked or bare; asm
Psilostacliyus, with naked spike.
Pteridium, Pterodium. Names for the
Key-fruit or Samara.
Pteris. Used for wing in Greek com-
pounds, also for a Fern.
Pteridof/raphia. The botany of Ferns.
Pterocarpous(-us). Wing-fruited.
Pteropodus. Wing-footed, i. e. petiole
wing-margined, &c.
Ptyxis. Greek name for folding, as of
leaves in a bud ; 132, 133.
Pubens, Pubfs Used for Pubescent.
Pub?rulus. Minutely pubescent.
Pubes. Pubescence, hairiness.
Pubescent (-ens). Clothed or furnished
with hairs or down, especially with soft
or downy and short hairs.
GLOSSARY.
Puf/ioniform (-ormis). Dagger-shaped.
Pullus. Dark-colored; dusky-brown or
blackish.
Pulvereus, Pulverulentus. Powdered ; as
if dusted with powdery matter or
minute grains.
Pulvinate (-atus), Pulviniform (-ormis).
Cushion-shaped.
Pulvinus. A cushion ; name given to an
enlargement or swelling close under
the insertion of a leaf, or sometimes
to the swollen base of a petiole.
Pumllus. Low or little.
Punctate (-atus). Dotted, either with
depressions like punctures, or trans-
lucent internal glands, or with colored
dots.
Puncticulate (-atus). Minutelj T punctate.
Punyent (-ens) Terminating in a rigid
and sharp point or acumination, like a
prickle.
Puniceous (-eus). Bright earn ine-red.
Purpurem. Originally the red of arte-
rial blood ; but our purple is some-
what dull red with a dash of blue or
violet.
Purpurdscens. Purplish.
Pusillus. Very small, or weak and slen-
der.
Pustular, Pustulate (-atus}, Pustulose
(-osus). Having low elevations, like
blisters.
Putamen. The shell of a nut; the endo-
carp of a stone-fruit; 288.
Pycnos, Greek for thick ; whence Pycno-
cephalus, thick-headed, &c.
Pi/ymceus. Dwarf, pygmv.
Pyramidal (-ali-<). Pyramid-shaped.
Pyrene (Pyrena). Same as Nucule or
Nutlet; one of the small stones of a
drupaceous fruit : 298.
Pyrtnarium, Pyridium. A pear or pear-
like fruit, same as Pomum.
Pyrenarius. Name of a drupaceous
pome, as of Medlar and Crata?gus.
Pyridion. Synonym of Pome.
Pyrenvcarp (-arpium). A general name
(or any drupaceous fruit ; 232.
Pijrifoi in (-ormis). , See Pear-shaped.
Pyxidntt, (-atus). Furnished with a lid.
Pijxidium, Pyxis. A capsule with trans-
verse dehiscence, making a lid of the
upper portion ; 293.
Quadri-. In Latin compounds, denotes
four; as Quadrangular, Quadri furi-
ous (in four vertical ranks), Quadriju-
gate (in four pairs), c.
Quaternary, Qnaternafe. In fours or
composed of four; 176.
Quini, Quinary (-ius), Quinate (-atus).
In fives ; 176.
Quinyue. Five. In Latin compounds,
giving rise to such terms as
Quincuncial, in a Quincunx; also five-
ranked ; 123, 136.
Quinquef'arious (-ius). In five vertical
ranks.
Quinquefoliate (-atus). Five-leaved.
Quinquefoliolate, with five leaflets.
Quintuple. Dividing into five parts, or
five-fold.
Quintuplinerred or -veined. With mid-
rib of leaf dividing into five (i. e. two
lateral pairs) above the base; 93.
Race. A variety of such fixity that it
is reproduced by seed ; also used in a
looser and more extended sense for a
series of related individuals without
particular regard to rank ; 320.
Raceme (Racemus). An indeterminate
or centripetal form of inflorescence
with lengthened axis and equal-pedi-
celled flowers; 146.
Racemif'erous. Bearing racemes.
Racemiform (-ormis). In the form of a
raceme.
Racemose (-osus). Having the character
or appearance of a raceme, or in ra-
cemes.
Rack is. See Rhachis.
Radial. Belonging to the ray.
Radiate (-atus). Spreading from or
arranged around a common centre,
or around the circumference of a cir-
cle ; bearing rays or ray-flowers.
Radiately veined. Same as Palmately
veined; 93.
Radiatiform (-ormis). Said of a capitu-
lum of flowers which is radiate by en-
largement of some of the outer flowers,
which however are not truly ligulate,
as in species of Centaurea.
Radical (-alis). Belonging to or pro-
ceeding from the root, or from a root-
like portion of stem at or below the
surface of the soil.
Radicant (Radicans). Rooting.
Radicel. A minute root or a rootlet.
Radiciflor, &c. In com-
mon English use, a male or staminate
flower is said to be a sterile flower.
Stichns. Greek for row or rank, usually
meaning vertical rank ; hence such
compounds as Distichous, two-ranked ;
Trfetichous, three-ranked, &c.
Stiyma, pi stigmata. That part or sur-
face of a pistil (usually on or a part of
the style, or in place of it) which re-
ceives the pollen for the fecundation
of the ovules ; 166.
Stiymatic (-tews), Stiymatose (-osus).
Relating to stigma.
Stiymatiferous. Stigma-bearing.
Stinys. Stinging hairs, seated on a
gland which secretes an acrid liquid,
as in Nettles.
Stipe ( Stipes}. A stalk of various sorts ;
the support of the cap of a mush-
room. ; the leafstalk of a Fern ; any
stalk-like support of a gynoecium or
a carpel; 212.
Stipel (Stipellum). An appendage to a
leaflet analogous to the stipule of a
leaf; 106.
Stipettate (-atus). Provided with stipels ;
106.
Stipitate (-atus). Having a stipe or
special stalk.
Stipitiform (-ormis). Shaped like a stipe;
stalk-like.
Stipulaceous (-eus), Stipular (-aris).
Belonging to stipules.
Stipulate. Possessing stipules.
Stipules. Appendages or adjuncts of a
leaf one on each side of the insertion ;
85, 105.
Stirps, pi. stirpes. A race.
Stock. Synonym of Race ; also the
portion of a stem to which a graft is
applied; a caudex, rhizoma, or root-
like base of a stem from which roots
proceed; 51.
Stole, Stolon ( Stolo). A sucker, runner,
or any basal branch which is disposed
to root ; 53.
Stoloniferous (-us). Sending of or propa-
gating by stolons, runners, &c.
Stoma, pi. stomata, Stomate. One of
the apertures in the epidermis of folia-
ceous parts, through which cavities
within communicate with the external
air; 89.
Stomatiferous (-us). Bearing stomata
or ''breathing pores."
Stone. The hard endocarp of a drupe.
Stone-fruit. A Drupe, such as a peach
or plum: 297.
Stool. The plant from which layers are
propagated, by bending down to the
ground to be rooted.
Stramineous (-eus). Straw-like or straw-
colored.
436
GLOSSARY.
Strap-shaped. See Ligulate (247) and
Lorate.
Striate (-atus). Marked with fine longi-
tudinal lines, streaks, or diminutive
grooves or ridges (Strice).
Strict (Strictus). Close or narrow and
upright; very straight.
Stritjillose (-osus). Minutely strigose.
Strigose (-osus). Beset with ftri^fe^ or
sharp-pointed and appressed straight
and stiff hairs or bristles.
Strobilaceotts (-eus), Strob'diform (-or-
mis). Relating to or resembling a
Strobile.
Strobile (Strobilus). An inflorescence
formed largely of imbricated scales,
as that of Hop and a Fir-cone ;
303.
Strombuliformif, Strombuliferm. Twist-
ed spirally into a screw shape, as the
legumes of the Screw-bean (Proso-
pis, sect. Strombocarpa) and of some
species of Medicago.
Strophiole (-tola). An appendage at
the hilum of certain seeds; 308.
Structural Botany, 2.
Struma. A wen or any cushion-like
swelling on an organ.
Strumose (-osus), Strumiferous (-us).
Furnished with a struma or goitre-like
swelling.
Stupose (-osus). Tow-like: with tufts or
mats of long hairs.
Style (Stylus). The usually attenuated
portion of a pistil or carpel between
the ovary and stigma; 166.
Styliform (-ormis). Style-shaped.
Styliferous. Style-bearing.
Stylinus. Belonging to the style.
Stylosus. With styles of remarkable
length or number, &c.
Styfopoflium. An enlargement or a disk-
like expansion at the base of a style,
as in Umbelliferae.
Sub. In composition of Latin words in
terminology, denotes somewhat or
slightly; as, Subacute,Subcordate,tha,t
is acutish, somewhat cordate, &c.
Subclass, 327.
Subconvolute and Subimbricate in aestiva-
tion, 137.
Suberose (-osus). Of a corky texture.
Submenus, 327.
Submerged, Submersed (-us). Growing
under water.
Suborder (Subordo), 327.
Subpetiolar (-aris). Under the petiole,
as the leaf-buds of Platan us , 42.
Subsection, 327.
Subspecies. A group which is ambigu-
ous in rank between variety and spe-
cies; 320.
Subtribe (Subtribus), 327.
Subulate (-afats), Subuli/brm (-ormis).
Awl-shaped.
Subvariety, 327.
Succise (-isus). As if cut or broken off
at the lower end.
Succubous (-us). When in leaves
crowded on a stem the apex of each
leaf is covered by the base of the
next above.
Succulent (Succosus). Juicv-
Sucker. A shoot of subterranean ori-
gin: 53.
Sujfrutesctnt (-ens). Slightly or ob-
scurely shrubby; 50.
Svjfrutex. An undershrub.
Sujfruticose (-osus). Low and shrubby
at base; 50.
Su/'ultus. Underpropped or supported.
Sulcate (-atus). Grooved or furrowed.
Super. Above. See Supra.
Superior, Superus. Growing or placed
above ; also in a lateral flower on the
side next the axis ; thus the poste-
rior or upper lip of a corolla is the
superior; 160, 183.
Superposed (Superpositus). Vertically
over some other part.
Superposition, 179, 195.
Supcrvolute (-us), Supervolutive (-ivus).
Same as Convolute when applied to
plaits; 139.
Supine (-inus). Lying flat with face up-
ward.
Suppression. Complete abortion; 179,
190.
Supra. Above; hence in Latin com-
pounds, Supra-axillary, above the
axil ; Suprafoliaceous,s\)ove. a leaf &c.
Supradedecompound. Several times com-
pound.
Surculose (-OSHS). Producing suckers.
Surculus. A sucker; a shoot rising from
a subterranean base ; 53.
Sursum. Upward; directed upward or
forward.
Su^ended (Suspensus). Hanging di-
rectly downward; hanging from the
apex of a cell.
Suspensor of the embryo, 284.
Sutural (-alis). Relating to a suture.
Suture (-ura). A junction or seam of
union ; used commonlj' as a line of
opening; 260.
Sword-shaped. A blade with two sharp
and nearly parallel edges, as in Iris.
GLOSSARY.
437
Syconium or Syconus. A multiple fruit
like that of the Fig; 148, 303.
Sylvestris. Growing in woods.
Symmetrical. Regular as to number of
parts or as to shape. In the blossom
it denotes the former; 175.
Symmetry. In the flower relates to
symmetrical disposition of organs on
the axis ; 174.
Sympetalous (-us). With united petals;
same as Gamopetalous; 244.
Sympkiantherous (-us). Same as Sy-
nantherous and Syngenesious.
Symphysis. Same as Coalescence.
Symphystemonous. With stamens united.
Sympode, Sympodium. A stem made
up of a series of superposed branches
in a way to imitate a simple axis; a
Sympodud stem ; 55, 154.
Synacmy. Same as feynanthesis.
Synantherous (-us). Stamens coalescent
by their anthers.
Synunthesis. The simultaneous anthe-
sis or readiness of the anthers and
stigmas of a blossom; 219.
Syncarp, Syncarpium. A multiple fruit
such as a mulberry, or a flesh}- aggre-
gate fruit, like that of Magnolia; 299.
Syncarpous (-us). Composed of two or
more united carpels; 261, 263.
Syncotyledonous. With cotyledons sold-
ered together.
Synedral. Growing on the angles.
Synema. The column of monadelphous
filaments, as in Mallow.
Synyenesia. Linnaean class (335) charac-
terized by having the anthers united or
Synyenesious. With anthers cohering
in a ring; 250.
Synonym. A superseded or unused
name ; 354, 365.
Synonymy All that relates to syno-
nyms ; 365.
Synsepalous (-us). Of coalescent sepals ;
same as Gamosepalous ; 244.
Systematic Botany, 2.
Systylus The coalescence of styles into
one body.
Tabescent (-ens). Wasting or shrivel-
ling.
Tail. Any long and slender terminal
prolongation.
Taper-pointed. See Acuminate.
Tap-root. A primary descending root
forming a direct continuation from the
radicle: 31.
Tawny. Same as Fulvous-, dull brown-
ish-vellow.
Taxoloyy, Taxonomy. Relating to clas-
sification and its rules ; 3, 315.
Teeth. Any small marginal lobes.
Teymen. The inner coat of a seed; 306.
Tela. Latin name for tissue, cellular
tissue, c.
Teleianthus. Same as perfect, or her-
maphrodite-flowered.
Tendril. A filiform production (either
axile or foliar) by which a plant may
climb; 54.
Tepal (Tepalum). A division of peri-
anth, whether sepal or petal (hardly
ever used).
Teratuloyical. Relating to malforma-
tion or monstrous conditions.
Teratology. The science of monsters
and malformations ; 170.
Terete (Teres). Round in the sense of
having a circular transverse section.
Teryeminate (-atus). Thrice twin.
Terminal (-alis). Proceeding from or
belonging to the end or apex ; 7.
Terminology. Same as Glossology; 3,
359.
Ternary (-arius). Same as Trimerous ;
consisting of three ; 176.
Ternate (Ternus, Ternatus). In threes;
as three in a whorl or cluster.
Tessellated (-atus). In chequer-work.
Testa. The outer seed-coat, which is
commonly hard and brittle, whence
the name, which answers to seed-shell ;
305.
Testaceous (-eus). Of the color of un-
glazed common (brownish-yellow)
pottery.
Tetra. In Greek compounds, four ;
hence
Tetracdrpellary (-arts). Of four car-
pels; 261.
Tefracdmarous (-s), Tetracoccus. Of
four closed carpels.
Tetradyndmia. Linnaean class (335)
which has the stamens.
Tetradynamous (-us). With four long
and two shorter stamens ; 250.
Tetrdf/onal or Tetrayonous (-us). Four-
angled.
Tetraijynia. Linnaean artificial order
(337), characterized by having the
gynoccium.
Teirdf/ynous. Of four carpels or styles.
Tetrdmerous (-us). Composed of four
members in a circle; 176.
Tetrandria. Linnaean class having the
flowers perfect and
Tetrandruus. With four stamens; 249,
334.
438
GLOSSARY.
Tetrapetalous (-its). With four petals;
244.
Tetraphyllous (-us). Four-leaved ; 243.
Tetraquetrous (-us). With four sharp
or salient angles.
Tetrasepalous (-us). With four sepals;
244.
Tetrdstichous (-us). In four vertical
ranks.
Thalamijtorous (-us), 340. With parts
of the flower hypogynous, or on the
Thdlamus. The receptacle of a flower;
167. See Torus.
Thallophytes ( Thallophyta), 341.
Thallus. A stratum, in place of stem
and foliage.
Theca. A case; an anther-cell (251);
a spore-case, &c. (An early name for
the anther, 166.)
Thecaphore (-orum). The stipe of a
carpel (homologous with petiole);
. 212.
Thorn. Same as spine ; 55.
Throat. The orifice of a gamopetalous
corolla or calyx, including any por-
tion between this and the proper tube ;
246. See Faux.
Thyrse, Thyrsus. A contracted or ovate
panicle; a mixed inflorescence, with
main axis indeterminate, but the sec-
ondary or ultimate clusters cymose ;
159.
Tiyelle, Tigellula. A miniature or ini-
tial stem ; sometimes applied to Cauli-
cle (Radicle), sometimes to Plumule;
10.
Tinctorius. Dyed; used for dyeing;
imparting color.
Tissue. The anatomical fabric.
Tomentose (-osus). Densely pubescent
with matted wool, or Tomentum.
Tonyue-shaped. Long and nearly flat,
somewhat fleshy, and rounded at the
apex.
Tooth. See Teeth.
Toothed. See Dentate.
Top-shaped. Inversely conical.
Torose (-osus). Cylindrical, with con-
tractions or bulges at intervals.
Tortuous (-osus). Bent or twisted in
different directions.
Torulose (-osus). Diminutively or
slightly torose.
Tortus. Twisted.
Tortilis. Susceptible of twisting.
Torus. The receptacle of a flower ; 167,
211.
Trabeculate (-atus). Cross-barred.
Trachea. A spiral vessel or duct, named
from resemblance to the tracheae of
insects.
Trachycarpous (-us) Rough-fruited.
Trachytpermous. Rough-seeded, &c.
Transverse (-ersus). Across; right and
left as to bract and axis; collateral;
160.
Trapeziform (-ormis), Trapezoid. Un-
symmetrically four-sided, like a tra-
pezium.
Tree. A woody plant with an elevated
trunk.
Tri-. In compound words, both Latin
and Greek, denotes three or triple.
Triachanium. A fruit like a cremocarp
but of three carpels.
Triadtlphous, Triadelphia. With fila-
ments in three sets; 250.
Triandria. Linnaean class (334) with the
flowers.
Tridndrous. With three stamens ; 249.
Trianyular(-aris), Trianyulatus. Three-
angled.
Tridnthous (-us). Three-flowered.
Tribe. Group superior to genus, in-
ferior to order ; 326.
Tricarpellary (-aris). Of three carpels;
261.
Tricarpous (-us). Consisting of three
fruits or carpels.
Tncephalous (-us). Bearing three heads.
Trichocarpous (-us). Hairy -fruited.
Trichodes. Resembling hair.
Tricholomous (-us). Three-forked;
branched into three divisions.
Trichome (Trichoma). Any outgrowth
of the epidermis, such as a hair or
bristle; 209.
Tricoccous (-us). Consisting of three
cocci.
Tricolor. Three-colored.
Tricuspidate (-atus). Tipped with three
cusps or pointed tips.
Tridentate (-atus). Three-toothed.
Tridlyitate. Thrice digitate.
Triduus. Lasting for three days.
Triennial (Triennis). Lasting for three
years.
Tiifarious (-ius). Facing three ways,
in three vertical ranks.
Trijid ( Tiijidus). Three-cleft.
Trifoliate (-ius, Trifoliatus). Three-
leaved.
Trifuliolate (-atus). Of three leaflets.
Trifurcate (-atus). Divided into three
forks or branches.
Triyamous (-us). Bearing three kinds
of flowers.
Trigonous (-us), Trigonal. Three-angled.
GLOSSARY.
439
Triyynia. Linnsean artificial order with
Triyynous, i. e. three-styled flowers ;
337.
Trihllatus. Having three apertures, as
in some grains of pollen.
Trijugate (Trijuyus). With three pairs
of leaflets or pinna;.
Trilobate ( Trilobus). Three-lobed.
Trilvcular (-aris). Three-celled.
Trimerous (-us). Three-membered parts
in threes; 170.
Trimestria. Lasting for or maturing in
three months.
Trimorphous, Trimorpism. Occurring
under three forms; 236.
Trinervate ( Trinervius). Three-nerved.
Trinodal. Of three nodes or joints.
Trioecia. Linnaan artificial order with
the flowers.
Tricecious or Trioicous (-us). Having
staminate, pistillate and perfect flowers
(or three kinds of flowers as to sex),
334; on three distinct plants.
Triovulate (-atits). Having three ovules.
Tripartible (-ibilis). Tending to split
into three portions.
Tripartite (-itus). Three-parted.
Tripetaloid (-oideus). As if three-pet-
alled.
Tripetalous (-us). Having three petals.
Triphyllous (-us). Three-leaved ; 243.
Tripinnate (-atus). Thrice pinnate; 104.
Tripinnatifid (-idits). Thrice pinnatifid.
Triple-ribbed or nerved. With midrib
dividing into three, or sending off on
each side a strong branch, above the
base of the blade ; 93.
Triplinerved (Triplinermus). Same as
Triple-nerved, Triple-ribbed ; 93.
Tripterous (-us). Three-winged.
Triquetrous (Triqueter). Three-edged;
with three salient angles.
Triquinate (-atus). Divided first into
three then into five.
Trisected (-its). Divided into three por-
tions; 99.
Trisepalous (-us). Of three sepals.
Triserial (-alls), Triseriate (-atus). In
three horizontal ranks or series.
Tristdchyus. Three-spiked.
Tristichous (-us). In three vertical
ranks; 122.
Tiistigmatic. With three stigmas.
Tristis. Dull colored.
Tristylous (-us). Having three styles.
Trisulcate (-atus). Three-grooved.
Triternate (atus). Thrice ternate; 104.
Trivial names, Nomina trivialia. Com-
mon or vulgar names; used by Lin-
naeus for specific names of a singlo
word ; 346, 362.
Trochlear (-earis). Pulley-shaped.
Trophosperm (Trophospermium). Name
for the Placenta; 261.
Trumpet-shaped. Tubular, with a dilat-
ed orifice.
Truncate (-atus). As if cut off at the
end; 97.
Trunk (Truncus). A main stem.
Tryma. A drupaceous nut, with exo-
carp at length dehiscent or otherwise
separating, such as walnut and hick-
ory nut.
Tubceformis. Trumpet-shaped.
Tube (Tabus). Any hollow elongated
body or part of an organ ; 245.
Tuber. A thickened and short subter-
ranean branch, beset with buds or
eyes; 59.
Tubercle (Tuberculum). A small tuber
or an excrescence: or something be-
tween a tuber and a root ; 60.
Tuberculate (-atus). Beset with knobby
projections or excrescences.
Tuberiferous. Bearing tubers.
Tubular, Tubulosus (-ose). Having a
tube; tube-shaped; 248.
Tubulijlorus (-us). When the flowers of
a head have only tubular corollas.
Tunicate (-us). Having coats (tunics).
Turbinate (-atus). Top-shaped.
Turion, ( Turio). A scaly sucker or
shoot from the ground ; 41.
Turnip-shaped. See Napiform.
Twin. In pairs. See Geminate, Didy-
mous.
Twining. Winding spirally and so
climbing ( Twiners); 51.
Twisted. Contorted.
Two-lipped. See Bilabiate.
Type. The ideal plan or pattern.
Typical. Representing the plan or
type.
Uliginose (-osus). Growing in swamps.
Ulndris. Of the length of the ulna or
fore-arm.
Umbel ( Umbella). An inflorescence
(proper!}' of the indeterminate type)
in which a cluster of pedicels spring
all from the same point, like rays of
f an umbrella; 146.
Umbellate (-atus), Umbelliform (-ormis).
In or like umbels.
Umbellet. A partial or secondary um-
bel ; 150.
Umbelliferous (-us). Bearing umbels.
440
GLOSSARY.
Umbellula. A partial or secondary um-
bel, or umbellet; 150.
Umbilicate (-atus). Depressed in the
centre, navel like.
Umbilicus. The hilum of a seed.
Umbonate (-atus). Bearing an Umbo
or loss in the centre
Umbrdculiform (-orrnis). Having the
general form of an umbrella.
Umbrosus. Growing in shady places.
Unarmed. Destitute of prickles, spines,
or other armature.
Uncate (-atus), Uncinate, (-atus), Unci-
form (-ormis). Hooked; bent or
curved at tip in the form of a hook.
Uncidlis. An inch (uncia) in length.
Undate (-atus) or Undulate (-atus).
Wavy; 98.
Undershrub. A very low shrub; 50.
Unequally pinnate. See Impari-pin-
nate.
Unyuiculate (-atus). Contracted at
base into an
Unyuis. A claw, or stalk-like base of a
petal, &c. ; 245.
Uni-. In Latin compound, one; as
Unictllular. Of one cell; Unicolor, of
f one color, &c.
Unicus. Singly or single, solitary.
Unijlorous (-us). One-flowered.
Unifoliate (-atus). One-leaved.
Unifoliulate, of one leaflet ; 102.
Unijuaate (Unijuyus). Of one pair;
102.'
Unilabiate (-atus). One-lipped, like the
corolla of Acanthus, in which the
upper lip is obsolete.
Unilateral (-alts). One-sided; either
originating on or more commonly
turned all to one side of an axis.
Unilocular (-am). One-celled.
Uninervate ( Uninervis, Uninervius).
One-nerved.
Uniovulate (-atus). Having only a soli-
tary ovule.
Uniparous. Bearing one: as a cyme of
one axis or branch; 152, 155.
Uniscrial (-ialis), Uniseriate (-atus).
In one horizontal row or series.
Unisexual (-alts, Unisexus). Of one
sex; having stamens only or pistils
^only; 191.
Univalved ( Univalvis). Of one piece or
valve.
Urceolate (-atus). Hollow and con-
tracted at or below the mouth, like an
urn or pitcher ( Urceolus).
Urens. Stinging, in the manner of net-
tles.
Utricle ( Utriculus). A small bladdery
pericarp; 295. Or any small bladder-
shaped body or appendage; also a
synonym of a cell of parenchyma.
Uiricular (-arts), Utriculate (-atus),
Utriculiform (-ormis), Utriculose
(-osus). Having or consisting of
utricles, or bladder-like in appear-
ance.
Vacillans. Swinging free, as the anth-
ers of Grasses on their filaments.
Vdcuus. Void or empty of the proper
contents.
Vayina. A sheath, as of a leaf, &c.
Vayinate. Sheathed.
V all-cuke. The intervals or grooves
betAveen the ridges or ribs of the fruit
Umbelliferee.
Valcate (-atus), Valvular (-arts).
Opening as if by doors or valves, as
do most dehiscent fruits (capsules),
and some anthers; also the parts of
a flower-bud when they exactly meet
without overlapping; 135.
Valve ( Valva). One of the pieces into
which a capsule splits, 288.
Valued. Same as valvate: hence 3
-valved, 5-valved, many valved, &c.
Vdlcula. A diminutive valve. Also
used (after Linnaeus) for the inner or
flower-glumes of Grasses.
Varieyated (-atus). Irregularly colored ;
in patches of color.
Variety ( Varietas). A sort or modifi-
cation subordinate to species; 318.
Variolate, Varioldris. Marked as if
by the pustules or pittings of small-
pox.
Vascular (-am). Relating to or fur-
nished with vessels ( Vasa) or ducts.
Vascular Plants ( Vasculares), 340.
Vdsculum. Same as Ascidium. Also
the botanists' collecting box ; 372.
Vasiform (-ormis). In the form of a
vessel, duct, &c.
Veined. Furnished or traversed with
n'bro-vascular bundles or threads, es-
pecially with, those which divide and
are reticulated.
Veins ( Vence). In general any ramifi-
cations or threads of fibro-vascular
tissue in a leaf or any flat organ;
especially (as distinguished from
nerves) those which divide or branch;
92.
Veinless. Destitute of veins.
Veinlet ( Venula). One of the ultimate
GLOSSARY.
441
or smaller ramifications of a vein or
rib; 93.
Vtlute (-atus). Veiled.
Vtliitinous (Velutinus). Velvety: the
surface covered with a soft coating of
fine and close silky pubescence, or
ve lumen.
Venation ( Venatio). The mode of vein-
ing; 90.
Venenatus, Venenvsus. Poisonous.
Venose (-osus) Veiny; abounding in
veins or network.
Ventral (-alis). Belonging to the an-
terior or inner face of a carpel, &c. ;
the opposite of dorsal.
Ventricose (-osus). Swelling unequally
or inflated on one side.
Ventriculose (-osus). Minutely ventri-
cose.
Venulose (-osus). Abounding with vein-
lets or venuhe.
Vermicular (-(iris) Worm-shaped.
Vernal ( Verndlis). A ppearing in spring.
Vernation (-iilio). The disposition of
parts in a leaf-bud; 132.
Vernicose (-osus). As if varnished.
Vermcose (-osus). Covered with warts
(verruca) or wart-like elevations.
Versatile ( Versutilis). Swinging to and
fro ; turning freely on its support ;
253.
Versicolor. Changing color, or of more
than one tint or color.
Vertex. The apex of an organ.
Vertical (-alis). Perpendicular to the
horizon ; longitudinal.
Verticil (-illus). A whorl; 6.
Verticillaster. A false whorl, composed
of a pair of opposite cymes; 159.
Verticillastrate. Bearing or arranged
in Verticillasters.
Verticillate (-ntus, -arts). Disposed in a
whorl; 6, 119, 120.
Vescide (-icula). A small bladder or
air-cavity.
Vesicular (-aris), Vesiculose (-osus). As
if composed of little bladders.
Vespertine ( Vespertinus). Appearing or
expanding in early evening.
Vessels ( Vasce). See Ducts.
Vexillary (-aris), Vexillar, 137. Per-
taining to the
Vexillum. The standard or large pos-
terior petal of a papilionaceous corolla;
184.
Villose (osus) or Villous. Bearing shaggy
or long and soft (not interwoven) hairs
or Villi.
Vimineous (-eus). Bearing long and flex-
ible twigs, like those used for wicker
work.
Vine. Any trailing or climbing stem :
originally that of the Grape from
which wine is made.
Vinedlis. Growing in vineyards.
Violaceous (-eus). Violet-colored.
Virens. Green, or evergreen.
Virescens. Greenish or turning green.
Viryate (-atus). Wand-shaped, or like
a rod ; slender, straight, and erect.
Vir(jultum. A vigorous twig or shoot.
Viridescent (-ens). Same as Virescens.
V it-id is. Green.
Viridulus. Greenish.
Virosus. Venomous.
Viscid (-idus)j Viscous (-osus). Sticky
from a tenacious coating or secretion.
Vitellinus. The yellow hue of the yolk
of egg.
Vitellus. Name formerly given to the
peculiar albumen which is in some
cases deposited within the embryo-sac.
Viticulose (-osus). Sarmentaceous ; pro-
ducing vine-like twigs or suckers,
viticulcB.
Vitt(z. The fillets or stripes (oil-tubes)
of the pericarp of most Umbellifera?,
which contain an aromatic or peculiar
secretion.
Vitiate (-atus). Bearing vittoe ; or with
any longitudinal siripes.
Viviparous (-us). Germinating or sprout-
ing from seed or bud while on the
parent plant.
Voluble. ( Volubilis). Twining round a
support; 51.
Volutus. Rolled np in any way.
Volva. A wrapper or external covering,
especially that of many Fungi.
Wavy. See Undulate.
Waxy. Resembling beeswax in consist-
ence or appearance.
Wedye-shuped or Wedge-form. See
Cuneate; 95.
Wheel-shaped. See Rotate.
Whorl. Arranged in a circle round an
axis; a Verticil; 6.
Wliorled. Disposed in whorls.
Wild. Growing without cultivation ;
spontaneous.
Winf/. See Ala. Any Membraneous
or thin expansion by which an organ
is bordered, surrounded, or otherwise
augmented. Also the two lateral
petals of a papilionaceous corolla are
termed wings; 185.
442
GLOSSARY.
Winged. See Alate : bearing a wing or
wings.
Withering. See Marcescent.
Wood. The hard part of a stem, &c.,
mainly composed of
Wood-cells, Woody Jibre or tissue, 68.
Woolly. See Lanate and Tomentose:
clothed with long and tortuous or
matted hairs.
Xanthos. Greek for yellow in com-
pounds, such as Xanthoplnjll, the yel-
low coloring matter in leaves.
Xenogamy. Fecundation of the ovules of
a flower by pollen from some other
plant of the same species; cross-fer-
tilization; 216.
Xylinus. Woody, pertaining to wood.
Zoospore. One of the free-moving spores
of the lower Cryptogams.
Zyfjomorphous (-us). That which can be
bisected in only one plane into similar
halves; 175.
ADDENDA.
Antidromous, Antidromy. When the course of a spiral is reversed, 157.
Infertile (-His). Said of a pistil or flower which fails to set fruit.
Polyembryony. The production of two or more embryos in a seed. 284.
Saprophytes (-yta). Plants feeding upon decaying vegetable or animal matter.
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