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3 (SloMBxz of ISotantcal Etxmsi. 


ASA GRAY, LL.D., etc., 


* .. I- ;• .' ;-. 

, V"'- * ^' 

• > ' . 





By Asa Gray. 


» <■ ^ w V 

*- * * ^ «F 


• • • 

• • 

1 494*52 

UinvERSiTT Press: John Wilson axd Son. 


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 Phaenogamous 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 sei-ve 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 Phoenogamous 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 
V^etable Physiology and Anatomy, and to the wide and 
varied department of Cryptogamic Botany. 


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 

A foui*th volume, a sketch of the Natural Orders of 
Phsenogamous 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. 


Herbarium of Harvard Univbrsitt, 
Cambridge, April 10, 1870. 

♦»♦ The numerals in parentheses, which are here and there introdnced 
into sentences or appended to them, are references to the numbered para- 
graphs in which the topic is treated or the term explained. 



INTRODUCTION. The Departments op the Science .... 1 





The Embryo, its Nature, Structure, and Parts 

Development of the Dicotyledonous Embryo in Maple ... 10 

In Ipomoea, or Morning Glory, &c., with Albuminous Seeds . 13 

In Embryos with thickened Cotyledons 16 

As of Almond, Beech, Bean, &c 17 

With Hypogteous Germination and no Elongation of Caulicle 19 

In Megarrhiza, &c., with concreted Petioles to the Cotyledons 21 
In Ipomcea leptophylla with foliaceous and long-petioled 

Cotyledons and no elongation of Caulicle 22 

In Pumpkin, &c., with no IMmary Root 22 

The Polycotyledonous Embryo 23 

The Monocotyledonous Embryo of Iris, Onion, Cereal Grains 24 

Pseudo-monocotyledonous and Acotyledonous Embryo ... 20 

Dicotyledonous and Monocotyledonous Plants 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 


Section II. Of Buds 40 

Scaly Buds and Bud-scales 40 

Naked, Subpetiolar, and Fleshy Buds 41 

Bud-propagatiun 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 Koot-clinibers .... 52 

Suckers, Stolons, Offsets, Runners 53 

Tendrils formed of Stems 54 

Synipodial and Monopodia! Stems 55 

Spines or Thorns and Subterranean Stems 50 

Rhizoma or Rootstock 57 

Tuber, Tubercles 50 

Corm or Solid Bulb 01 

6ulb, Bulblcts 02 

Condensed Aerial Stems 04 

Stems serving for Foliage, Phyllocladia, Cladophylla ... 05 

Frondose Stems 60 

§ 3. Internal Structure 07 

Anatomical Elements 08 

Endogenous Structure 70 

Exogenous Structure ; its Beginning 73 

First Year's Growth 74 

Pith, Layer of Wood, &c 75 

Bark, its Parts and Structure 70 

Annual Increase in Diameter 78 

Demarcation of Annual layers 70 

Sap-wood an<l 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 T^eaf 85 

Duration, Defoliation, Normal Position 80 


§ 2. Their Structure and Forms as Foliage 87 

Internal Structure or Anatomy 87 

Parencliyma-celU 88 

Epidermis, Stomata or Breatliing-pores 89 

Framework, Venation 00 

Parallel-veined or Nerved Leaves 91 

Reticulated or Netted-veined Leaves 02 

Pinnately or Feather-veined and Palmately or Hadiately 

Veined r3 

Forms as to Outline 94 

Forms as to Extremity 00 

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 100 

Such as Insquilateral, Connate, Perfoliate 107 

Vertical and Equitant 108 

Without distinction of Parts 109 

Stipules serving for Blade 109 

Phyllodia, or Petioles serving for Blade 1 10 

§ 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 110 


Section I. Distribution of Leaves on the Stem .... 119 

Pliyllotaxy either Verticillate or Alternate, Cyclical or Spiral 119 

Verticillate or Cyclical Arrangement 120 

Alternate or Spiral Arrangement 121 

Its Modes and I>aws 122 

Relation of Whorls to Spirals 129 

Hypothesis of the Origin of Both \m 

Fascicled Leaves 131 

Section II. Disposition of Leaves in the Bud 132 

Vernation and Estivation; the Modes 132 

Direction, Dextrorse and Sinistrorse 140 

• • • 



Bracts and Bractlets and their Modifications 141 

Peduncles, Pedicels, Rhachis, Receptacle 143 

Position of Flower-buds, Kinds of Inflorescence 144 

Lidetenninate, Indefinite, or Botryosc 14C 

Raceme, Corymb, Umbel 140 

Head or Capitulum 147 

Syconfum 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 Heliooid, the Pleiochasium, Dichasiom, and 

« Monochasium 155 

Bostryx, Cincinnus, Rhipidium, Drepanium, &c 150 

Mixed Inflorescence 158 

Thyrsus, Verticillaster, &c 159 

Relations of Bract, Bractlet, and Flower 100 

Anterior and Posterior, or Inferior and Superior 100 

Median and Transverse 100 

Position of Bractlets 101 

Tabular View of Inflorescence 102 


Section I. Its Nature, Parts, and Metamorpht .... 103 

Floral Envelopes, Perianth, or Perigone 104 

The Parts, Calyx and Corolla 105 

Androecium, Stamens 105 

Gynoecium, Pistils 100 

Torus or Receptacle of the Flower 107 

Metamorphosis 107 

Unity of Type illustrated by Position and Transitions . . . 109 

. Teratological Transitions and Changes 170 

Section II. Floral Symmetry 174 

Symmetrical, Regular, and Complete Flower 175 

Numerical Ground-plan 170 

Pattern Flowers 170 

Diplostemonous Type 177 

Section III. Various Modifications of the Flower . . . 179 

1. Enumeration of the Kinds 179 

2. Regular Union of Similar Parts 180 

Coalescence or Cohesion 180 


§ 3. Union of Dissimilar or Successive Pabts .... 181 

Adnation or Connation 182 

Hypogynous, Perigynous, Epigyuoiu 183 

§ 4. Irregularity of Similar Parts 184 

§ & 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 AndrtBcium 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 190 

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, Car]X)phore, &c 212 

Disk 213 

Hypanthium 214 

Section IV. Adaptations of the Flower to the Act of 

Fertilization 215 

§ 1. In General 215 

Close and Cross Fertilization, or Autogamy and Allogamy . 210 

§ 2. Adaptations for Allogamy or Intercrossing . . . 216 

Wind-fertilizable or Anemophilous Flowers 217 

Insect-fertilizable or Entomopliilous Flowers 218 

Irregularity as related to Allogamy 219 


Dichogamy, either Proterandrous or ProterogTOons .... 210 

Proterogyny 219 

Proterandry ; 220 

Particular Adaptations in Papilionaceous Flowers .... 225 

In Kalmia-blossoms, Iris, &c 229 

Transportation of PoUinia 230 

In Orchidaceae and Asclepiadacee 231 

Heterogonous Dimorphism and Trimorphism 234 

§ 3. Adaptations for Close Ferti uz ation 240 

Cleistogamy 241 

Section V. The Perianth, or the Calyx and Corolla in 


Perianth as to Duration, Numerical Terms, Union, &c. . . . 243 

Parts of Petals and of Gamophyllous Perianth 246 

Forms of Corolla and Calyx 240 

Section VI. The Andr<ecium, 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 YU. The Pistils, or Gyncecium 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 PlacenU 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 Cycadaceaa 274 

Section VIII. The Ovule 276 

Its Stnicturc and Position 277 

Its Forms, Orthotropous, Campylotropous, Amphitropous, 

Anatropous 278 

Origin and Morphological Nature of the Ovule 282 

Origination of the Embryo 283 



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 Antliocarpous Fruits 300 

Multiple or Collective Fruits, Syconium, Strobile, &c. . . . 801 

Table of Simple Fruits 804 


Its Stalk, Coats, and Appendn^s 306 

ArilorArillus 308 

Nucleus or Kernel, Albumen 309 

The Embryo, its Parts and Positions 31 1 

Tlie Cotyledons as to Adjustii.ent and Number 313 


Section I. The Principles op 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 

Antc-Linnsan Classifications 332 

Liniuean Classification 333 

Sexual Artificial System 3:U 

Natural System 338 

As presented by Jussieu 339 

Some of its Modifications 340 

• • 



Section I. Nomenclature 345 

Names of Plants, Binomial Nomenclatore 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 350 

Tribal and Ordinal Names 357 

Names of Cohorts, Classes, &c 858 

Section II. Glossology or Terminology 851) 

Section III. Description 361 

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 

Tlie Herbarium 380 


SIGNS 391 






1. The two Biological Sciences,^ considered as parts of Natural 
Histor}', are Zoology and Botany. The latter is the natural 
histor}' 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 econom}' of the world. 

2. We cannot distinguish the vegetable from the animal king- 
dom by any complete and precise definition. Although ordinary 
obser\'ation 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 j)lants 
ix>sscssing 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 stnicture, and of which the bodies 

* Bioltyjif, 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, i*. e. of the two organic kingdoms, including 
both their physiology and descriptive natural history. 



of plants and animals are composed, is called organic matter. 
Animals appropriate and live u^wn this, but have not the ix)wer 
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, yd plants also consume organic matter, 
more or less, acting in this resi)ect like animals in all their opera- 
tions, except in the grand and peculiar one by which tliej' 
assimilate mineral matter. Most plants of the higher grades 
assimilate largely and consume Httle, except in special oi>era- 
tions. Some, on the contrary, are mainly consumers, and feed 
iil)on foimed organic matter, living in this resixjct 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 Physiology 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 Ph^'siologj'. 

5. STRUCTrRAL BoTANY comprehcnds 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 t^-pe. The terra 
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 microscoiK?, t. 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 stnicture, and 
indeed answers to organography, 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 derive<l IVom the facts and ideas 


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 Nomenxlature, 
the methods and rules adopted for the formation of botanical 
names. Glossology or Terminology^ is a necessary part of 
Photography 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, by the use of 
which the botanist is able to describe the objects of his study 
with a precision and bre^ity 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 api^nd to this 
volume an alphabetical summarj- of them, or Vocabularj' of 
Botanical Terms.* 

7. The present volume is mainlj' devoted to Morphological 
Botan}' ; that is, to Structural Botany on the basis of mor- 
pholog}'. This department cannot be properly dealt with apart 
from considerable reference to intimate stnicture, development, 
and function, the subject-matter of vegetable histology and 
pho'siologj-. But these will here be treated only in the most 
general or incidental and elementarj' way, and only so far as 
is necessary Uf the understanding of the morphology' of the 
stem, leaves, '&c. The whole discussion of the histology and 
plnsiolog}' 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, t. e. into PHiENOGAMOus (or Phanerogamous) or 
Flowering, and Cryptogamous or Flowerless Plants. The 
first are all manifestly of one to'pe, and therefore have a consist- 
ent and simple moq^hology. The second differ among them- 
selves almost as widely as they do from the higher series ; and 

1 Glossoloot is the better word, but Terminology, although a hybrid 
of T^tin and Greek, is in common use. 

* 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 caases 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. 


their morpholog}' is more special and difficult. Wherefore it is 
better to treat them separatel}' and subsequently. This will be 
done in a third part, by an associate devoted to Cryptogamie 

9. Thus the field is here left clear for the Structural Botany 
of Phflenogamous or Flowering Plants, with which the study of 
the science should naturally' begin. In theory" it may seem 
proper to commence witli the simplest plants and the most ele- 
mentary structures ; but that is to put the difficult and recondite 
before the plain and obvious. The tyi)e or plan of the vegetable 
kingdom, upon which morphological botany is grounded, is fully 
exemplified onl}- in the higher grade of plants, is manifest to 
simple obser\'ation, and should be clearly apprehended at the 



10. Morphology, the doctrine of forms, as the name denotes, 
is used in natural histor}- in nearly the same sense as the older 
term Comparative Anatomy. If it were concerned merely with 
the description and classification of shajxjs and modifications, 
it would amount to little more than glossology and organography'. 
But it deals with these from a peculiar point of view, and under 
the idea of unity of plan or type.* 

11 . As all vertebrate animals are const ructe<I 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 t^pe ; 
the different kinds l>eing patterns or i-epetitions of it, with varia- 
tions. The vegetable kingdom, however, does not culminate in 
an archet>i^e or highest i*epresentative. As resi>ects 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 crj'ptogamous plants are left out of the 
present view, to be treated apart. 

12. Viewed moq)hologically and as to its component oi'gans, 
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 verj' unlike leaves in whole appearance 

* The term Morpholwjy was introdnce<l into science by Goethe, at least as 
early as the year 1817 (Zur Naturwissenschaft iiberhaupt, besontlers zur 
Morphologie, Stuttgart und Tiibingen, 1817-21). On page 9 of the first 
volume, he is understood to have suggested this word for tlie purpose and in 
the sense now adopted in botany and zoology. It essentially replaces an 
earlier and somewbat misleading word, Mitmnm-pttosis. (IJOl.) 

Apparently the first botanist to adopt tbe term was Auguste dc St. 
Hilaire, in his "Lemons de Botanique, comprenant principalement la Mor- 
phologie Veg<?tale, etc., Paris, 1841. Tbe term seems not to have been taken 
up, in zoology, by Etienne Geofifroy Saint-Iiilaire, the antagonist of Cuvier 
(who was of a wholly different family from tbat of the botanist), although 
tbe same idea was denoted by his phrase " miity of organic composition." 


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 ordinar}' form is 
the familiar one of foUage ; hence the name of leaves has been 
by botanists extended in a generic wa}' from the green expan- 
sions which constitute foliage to other forms under which such 
appendages occur. The proper morphological expression is, 
that the latter ai*e homologous with leaves, or are the homologues 
of leaves.^ 

13. Leaves are borne upon the stem at definite places, which 
are terme<l Nodes. A node may bear a single leaf or a greater 
number. AVhen it bears two, tliey 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 onl}- 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 cdter- 
nate^ that is, come one after another on the stem ; or in whorls 
{whorled, verticiUate) ^ in the commoner case of a single pair 
being called opposite. The bare space between two successive 
nodes is an Internode. This is longer or shorter, accoixiing to 
the amount of longitudinal growth, which thus spaces the leaves, 
or whorls of leaves, in most various degrees, either widely when 
the intemodes are elongated, or slightly when they remain very 
short. The plant, therefore (roots excepteil), is made up of a 
series of similar parts, i, e, of portions of stem, definitely bearing 
leaves, each portion develoi>ed 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 fonns with the stem, — 
from which they grow much as the prima r}' stem grew from the 
seed. The primary stem, connected with the ground, produces 
roots which develop downwanll}' into the soil, from which they 
draw sustenance. Branches, when develoi>ed above ground, 

' A common designation for all these appendages being desirable, a good 
one is furnished by the Greek name for leaf, ^vWov, Phylum, plural 
Phylla. This, used with prefixes, may be made to designate the kind of 
leaves in many cases, — as, pvphylia^ rnta/thy//a, hypsophfiUa. 

Recent German botanists use the word Phylhme in this sense. It is a 
rather convenient and well-sounding word ; but phylloma 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 phylinm, readily take prefixes, as above, or the adjec- 
tive form, as it readily does in pro^fhylhus, hypaophylious, gamophyllous, &c. 


being in organic connection with their parent stem, do not 
usaally produce roots; but when placed in equally favorable 
conditions for it, t. 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 nidimentary 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.^ 

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 progcn}', 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 develoi>ed, from ax- 
iUaiy buds. These ultimate similar parts into 
which a plant may thus be anal3'zed, and 
which are endowed with or may produce all 
the fundamental organs of vegetation, were by 
Gaudichaud called Piiytons. 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 (qpi'Tor, plant, 
^/(><V, part) , equivalent to plant-parts, — the 
structures which, produced in a scries, make 
up a plant of the higher grade. In English, 
the singular may be shortened to Piiytomer. 

1 7. This theoretical conception of the organic 
composition of the plant is practically impor- 
tant to the correct understanding of morpho- 
logical botany. The diagram, Fig. 1, serves 
to represent the organic elements, or phytomera, 
in a simple case, such as that of a growing 
plant of Indian Corn, or other Grass. Here 

1 Bifurcation by the division of a terminal bud into two, ns in Acrojycnous 
Cryptogams, is supposed by some to occur, even normally, in some Phwno- 
gams, especially in certain forms of inflorescence ; but this lias never been 
convincingly made out. 

FIQ. 1. Diagram of a 8lmple-»teniincHl plant, exhibiting tho niniilar parts, or 
phytomera, a to A, of which it is composoU. 


the leaves are alternate; in other words, each ph3'tomer is 
single-leaved ; while in the subsequent illustrations of plants 
developed from the seed, at least the earliest phj'tomera are 

18. The plan thus exhibited in the leaf}- stem begins in the 
embiyo, or initial plant in the seed, and is carried on into the 
flower, in which the normal development of the axis finall}' ends. 
One plan prevails throughout. To illustrate it, the moqjholog}* 
and growth of the embryo, of the plant developed for vegetation 
and the general purposes of its individual existence, and lastly 
of the flower, through which sexual reproduction takes place, 
may be successively treated in this order. 





19. The Embrjo is the initial plant, originated in the seed.^ 
In some seeds it is so simple and rudimentaiy as to have no 
visible distinction of parts: in others, these parts may have 
assumed forms which disguise their proper character. But ever}'^ 
well-developed embrj'o essentiall}' consists of a nascent axis, or 
stem, bearing at one end a nascent leaf or leaves, or what an- 
swers to these, while from the other and naked end a root is 
normall}' to be produced. This stem is the primitive inteniode 
of the i)lant : its leaf or pair of leaves is that of the first node. 
The plant therefore begins as a single ph^-tomer. Some embryos 
are no more than this, even when the}* have completed their 
proper germination : others have taken a further development 
in the see<l itself, and exhibit the nidimcnts of one or more fol- 
lowing phytomera. The embryo of the Maple is an example of 
the first kind ; and, being lai-ge enough for handling and for the 
display of all its parts to the naked eye, and the character of 
these parts being manifest even in the seed, it is a good subject 
with which to commence this study. And for this the Sugar- 
Maple is one of the best of 
the Maples. Its embr^'o 
(seen in Fig. 2 in the coiled 
condition which it occupies 
in the seed, and in Fig. 3 
and Fig. 4 uncoiling and be- 
ginning to grow) is an initial stem, bearing a pair of leaves, and 
nothing more. These parts take the technical names of 

1 Normally a send contains a single embryo. P(^i/emhrff, tlie format ion of 
two or more embryos, occurs occasionally as a kind of supcrfirtation in 
some seeds. In those of the cultivated Orange it is most common, and an 
evident monstrosity. In Coniferae and Lorantliacea;, two or three embryos, 
of equal size and perfection, arc not rarely produced. 

FIG. 2. Embryo of Sns^r Maple, in vertical section, as coiled in the 8ee<1, merely 
somewhat loosenefl. 3. Embryo of game, Junt l>eginnin(; to unfolii in germination. 
4. Same more advanced : a. ita stem or caulicle ; bb. its two leaves or cotyledons. 



20. Canlicle or Radicle, and Cotyledons. The name of radicle 
was earl}' applied to the axis of the embr3'0 below the cotyledons, 
on the supposition that it was the actual beginning of the root. 
But its structure and mode of growth show it is not roQ t-(24, 
44, 78), buLJLJ>o<b'-.Qf the exact natui:e_of _stei», f rom the 
u^aked.ejid of whicii the root is developed. Wherefoi-e CaulicU 

(Lat. cauliculus^ diminutive of caulis, stem) is 
the appropriate name ; and it would be gen- 
erall}' adopted, were it not that the older term 
is so incorporated into the language of sys- 
tematic botany (in which fixity and unifonnity 
are of the utmost importance) that it is not 
easily displaced. It may be continued in 
descriptive botany on this account, but in 
morpliology it is apt to mislead ; and the name 
of canlicle^ suggestive of the true nature of 
the organ, is prcfemble.^ The more fanciful 
name of Cotyledons was very early applied to 
what are now recognized as answering to the 
leaves of the embryo : it has the negative ment 
of suggesting no misleading analogy.* 

21. Development of the Dicotyledonous Em- 
bryoy i. e. the two-leaved embrj'o. This, in 
the Red Maple (Figs. 5-8), usually germinates 
in summer, shortl}' after the fruits of the season 
have matured and fallen to the ground. It 
differs from that of Sugar Maple in the crump- 
ling instead of coiling of the cotyledons in the 
seed. Referring the whole physiologj' of ger- 
mination to that part of the work which treats 
of Vegetable Physiolog}', the development of 
(he embr3'o into the seedling may here be described, taking that of 
a Maple for a convenient i\\^ or pattern, with which other forms 

* Linnaeus called it liostillum, a name wliicli, being etymologically mean- 
ingless in this connection, is not misleading. The French botanists named it 
Tigeile, diminutive of tige, stem : but some (like Mirbel) applied the term to 
the developing axis above the cotyledons; others, to the early axis both 
above and below them. The name ItfuUmUi originated with Gaertner. 

2 The name Cdyledon, which was adopted by Linnaeus, is a Greek word 
for a cup-shaped hollow or cavity, also for a plant with thickish and saucer- 
shaped leaves. It was primarily applied to the thickened " lobes " of the 
embryo, the foliaceous nature of which was not recognized. 

FIG. 5. One of the twin winjsred fruits of Re<l Maple (Acer rnbrnm), with ho«ly 
divided, to show the need. 6. See<l cxtrarteil and i!i\ ((1(m1. to show the embryo within. 
7. Embryo partly unfoldeil. 8. Embryo in early Ptago of germination. 



ma}' afterward be compared. The first growth is seen in the 
elongation of the radicle or caulicle, and its assumption, as far as 
possible, of a vertical position, and the production of a root from 
the naked end. As it emerges from the seed in consequence of 
this elongation, the root-end of the caulicle points downward into 
the soil, the caulicle bending, if need be, to assume this position ; 
and the nascent root, partaking of this disposition, grows in a 
downward direction. Hence the root has been called the Descend- 
ing Axis of the plant. While this avoids, the opposite or budding 
end (as it may be termed) seeks the light, and when free takes 
an upward direction. The result of this, and of the elongation 
of the caulicle, is to carrj- the budding end out of the soil and 
into the air, where the growing cot3'ledons unfold or expand and 
become the first leaves, or Seed-leaves, This initial stem and its 
continuation therefore constitutes the Ascending Axis, If the 
budding end happen to Ue pointing downward and the root-end 
upward in the ground when germination begins, both will cun'e 
quite round, as they grow, to assume their appropriate directions. 
If obstacles intervene, each will take as nearly as possible its 
wonted direction, through an instinctive tendenc}" and action, 
which insures that each part of the plant shall be developed in 
its fit medium, — the root in the dark and moist earth, the stem 
and leaves in the light and air. 

22. The plantlet, thus established, has now all the essential 
Organs of Vegetation^ as the}' are called, 
». e, root, stem, and leaves. Its subse- 
quent development, so far as vegetation 
(apart from proper reproduction) is con- 
cerned, consists in the addition of more 
of these, until the whole herb, shrub, or 
tree is built up. 

23. In Maples (as in the Morning Glory, 
Fig. 16, and many others) the embrj'o in 
the seed, and until after the full develop- 
ment of its cotyledons or seed-leaves, 
shows no rudiments of the subsequent 
growth. The embrj'o grows into the plant- 
let wholly b}' the appropriation of prepared 
nourishing matter which was provided b}' 
the mother-plant and store<l in the seed, — in the case of the 
Maple, wholly in the embiyo itself, mainly in its cotyledons. 

FIQ. 9. Maple embryo developed into plantlet of one phytomer, and pro<lucing 
Ttulimentfl of the second: the lower portion covered with root-holrs is the root; the 
naked portion above is the cauUcle. 



After this is consumed and in good part conveiled into struc- 
ture, the plautlet must by the action of its root and loaves imbibe 

from the soil and air appropriate materials, and 
assimilate them into nourishing matter needful 
for further growth. Onl}' then does the rudi- 
ment of new structure appear, in the form of a 
growing point, or bud, at the node or a\yex 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 tiie second node develops in 
the same wa}' 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 diflerent mode. 
The primordial stem, pre-existing in the seed 
(though at first it maj' be extremel}' short) 
grows throughout its whole length, but most 
in its upper part, so that it may Ixjcome a 
stemlet two or three inches long. But, soon 
attaining its full growth as to length, the 
stem is carried upwards b}* the subsequent joints or portions, 
similarly developed and elongated, one after the other. >^'ot 
that each ix)rtion necessarih' 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, — j'et 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 ix)tentialh' in tissue of the caulicle 
fix)m which a root or roots normall}' originate.^ It is fonned 

^ Yet from nothing which is special to this part of the embryo, nor to the 
embryo at aU. Tlie 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. tlieir 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 and pair of loaves complete, and bud of the third apparent. 




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 
ca{) or sheath), b called the Root-cap, ^ 
As the primal^- 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 tlie old. That is, the 
root elongates by continual minute 
increment of its apex or near it, the 
formed parts very soon ceasing to 
lengUien. This is in marked distinc- 
tion from stem, which grows by suc- 
cessive individualized portions ; and 
these portions (intemodes) , 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. 
They commonly branch or divide, but 
in a vague manner ; and their new parts bear what are called 
Jiool-hairs, which greatly increase the absorbing surface ; other- 
wise they are destitute of appendages or oi^ans. 

25. With the Maple embrjo, here taken as a ty|», that of 
Morning Glory, Ipomtca puri»urea, or any of its kin, may next 
be compared. The cotyledons are different in shape, being as 
broad as long, and notched both at base and a\wx. They lie 
in contact in Fig. 14, and are vcrj- 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 embrjo, as it lies in 
the seed, exactly di^'ides them (passing through the tenninal ami 
basal notches) and also the caulicle. which here is thicker tlian 
both. The germination is similar to that of the Staple ; and like 
that (as Fig. IG shows), and for the same reason, no bud or 
rudiment of the (brther growth pre-exists in the embryo or 

liree Julntsof Item an J pnjnoriea 




Appears in the joung plantlet, until that has established itaelf 
and had time to chiborate proper mati-rial therel'ov. Tliis wjn- 
(lition 18 conx'lated with thin foliaeeous 
cotyledons, holding no stiirc of nourieh- 
f iu«nt. Here they do not contain eiiiUciont 
material for the development of the initial 
flt*-m and root. The maternal p^o^■isioI) 
for ttiia is here stored up in th« seed 
round but not within the cnibrjo. This 
nourisliiiig deposit, seen in tlie section 
) (Kig. 13) filling the whole eiuice Iwtween 
the seed-eoate and tlie thin embryo, was 
named by the early botanists and vege- 
table auatomista the Albcheh of the seed.' 
This Bubstanee. softened in germination 
and by ehemical changes renderetl soluble, 
ia gradually absorbed by tlie cotj'ledons 
as malarial for tlioir growth and that of 
the developing primary stem and root. 

20. Seetls in this regard are aecortlingly 
distinguislied into albtiminoiu and rral- 
buminoui, those supplied with and those 
destitute of albumen. The difference 
" " inheres neither in the character nor in 

the amonnt of the maternal pro^^Bion for the development of 
tlie einbryo-|ilniit. but merely in the storage. In exalbuminous 
seeds the nourishment supplied for this purpose is taken into 
the euitiryo itself, mostly into the cotyledons, iluriag the growth 
and W-fon.' the maturity of the see<l. In alliuminoua seeds 
tills same material ia deix)sited around or at least external to 
the embrjo. 

27. Tlie amount of this deposit is, in the main, inverse)}' pro- 

' Grew appMira to \iaxm Hnt applied Ihii nAme, ami GHertmr to Imve 
inlrodun^ il inlo systcmaiic Itoiany. where it remains in use. aitlioueh 
Juuleu replm^wl it by the Ipnn IWUperui, and Itieharit liy EndotiirTm, 
neither of dipni much better etymolotiieally than the old word AllniBiiin. 
But il muKl W kept in mind that il was Intended in liken tlie " albumen " o( 
tlie lecil willi tlie albumen or wliitc of an viig "* *> ^x"'/ or niaM, and not ai 
a ehemlcal aulHiancc: the embryo being lani'irutly timceireil Id lie aiialo- 
gou) to the y«U- of the c'gg. the iiirruuniling lubatanov u( llii* kind nut 
uniutarallj' rook tlie nami> of the tehltt, t'a. albumen. 

PIQ. 13. Scrilim nt hwI at rmman Momlni Qlnrr. Ipmnoia parpuna. [lltlilJiii 
tliiionnulimlFintirfncliAiutlitbceBiiln, II, Knibryfi nf utne, iIhIacIib-I ami ninlclir. 
cnol. IG, EiDbryolnfnmliiaUini: UieEnl;1ei)<imuiilT jiaRl)' daUcIinl IVnni tliu i"iaC 
at ihK wnt. Irt. Same. Ini«r and mora iterdoiwd, ths aMjrlixlani unf»l>lnl aiiil ont- 
•■•raail u (ha Dm pair at luaria. 





portional to the size and strength of the embryo, 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 upper rank of 
figures represents 
sections of seeds ; 
the embr3*o left in 
white ; the albumen 
as a dotted surface. 
The lower rank shows the embrj'os detached. That of Mirabilis 
has verj' broad and thin cotyledons, 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 cotj'ledons narrowed down to 
semi-cylindrical bodies, not leaf-like in appearance, and the two 
together not thicker than the caulicle. In Barberr}- the embr3-o 
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 Peon}- 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 di\'ided, that is, to consist of two minute cotyledons. 
The embryo 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 the}' develop 
in germination. 

28. The stud}' of the formation of the embr3'o in the seed 
teaches that all embrjos 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 embryo 
while yet in the seed. It also appears that the size and shape 

FIQ. 17. Section of seed aiMl contained embryo of Mirabilis (Fuar-o-clock). 
Id. Embryo detached entire. 

FIG. 19. Section of a Potato-need. 20. Embryo detached entire. 
FIO. 2L SecUon of BarlMrry-need. 22. Embryo detarhed enUre. 
FIQ. 23. Section of Peony-eeed. 24. Embryo detached enUre. 



of an organ do not indicate its nature, either in the embrj'o or 
in subsequent growth. But in all the cases yet mentioned the 

cotyledons actuall}' demonstrate their 
nature by developing in gennination 
in a foliaceous manner and becoming 
the first leaves of the seedling. Nor 
is this nature much disguised by the 
fact that the}' differ greatly in form in 
different species, and that the seed- 
leaves, or developed cotyledons, differ 
much in shai)e 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 which previous to germination have 
developed nothing be3ond the cot3'ledons. 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- 

FIQ. 25. Further development of Morning Glory, Fig. 16. the root cut away, the 
internode above tlie cotyle<lon8 and its leaf complete*!, the next internode and Its leaf 

Flo. 2G. Embryo (kernel) of the Almond. 27. Same, with one cotyledon rcmovefl, 
to show the plumule, a, 

FIG. 2«. Section of an Apple-9ec<l, magnified, cutting through the thickness of the 
cotyleilons. 211. Embryo of the same, extract^yl entire, the cotylwlons a little separate*!. 

FIG. 30. Germination of the Cherry, showing the thick cotyledons little altered, 
ftnd the plamule developing the earliest real foliage. 



dons, t. e. cotyledons well charged with nourishing matter. The 
earl}' vegetable physiologists gave to it the name of Plumule 
(Lat. plumulay a little plume). The 
name was suggested by its appearance 
in such an embiyo as that of the bean 
(Phaseolus) , in which it evidently con- 
sists 9f a loidimentarj- pair of leaves, 
while in the pea and the acorn it is a 
rudimentar}' 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 embr}-o, 
that of an apple-seed, enlarged to 
nearly the size of the other. It is 
equally visible in the cheny, the bean, 
and the beechnut. The embiyo in all 
these cases constitutes the whole kernel 
of the seed. For the nourishment, 
which in all the foregoing illustrations 
except the first (».«. in Fig. 13, 17-23), 
is deposited around or exterior to the 
embrj'o, is in these stored within it. 

31. The development of these em- 
br}'08 in germination proceeds in the 
normal manner, but with two cor- 
related peculiarities. First, by the 
lengthening of the radicle more or less, their thick cotyledons 
are usually raised to or above the surface of the soil ; the}' 
expand, assume the green color needful to foliage ; but they 
imperfectl}' 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 IkjIow 
and the ready-formed plumule al3ove grow rapidly and strongl}', 
having accumulated capital to draw upon ; and the leaves of the 

FIG. 31. Beecbnut cat acro«s, filled by the flishy embryo; the thick cotyloilons 
partly enfolding each other. 32. Embryo of the Mvtiio in early germinal ion 33. Same 
more advanced; the plamule, which is Jnit emerging in the precc<ling, here developed 
into a long intemode and a pair of leaves. 




latter are practically the earliest efBcient foliagic of the plantlet. 
Thus, as ill the gcrmiiinting Cherrj--see<l (Fig. 30) , three or four 
intcmottes of stem, with their leaves, may bo produced before 
these leaves tiiemaelvos are Bulliciently fleveloi>ed to make any 
sensible contribution to this growth. And in the Beech and Bean, 

the leaves of the plumule come forward almost lieforo the root 
has attacliod the plantlct to the soil, (Fig. '62. 3.').) Between 
such cases and that of Maple and liie hke there are all degri-crt. 
There an.^ also familiar casea in which the storage of noui-ishmeiit 
in the cotylwiims is carrieil to a muxiinum, with results which 
gravely afFect the doveloitment. 


31. Tlia ftti 

lirjn (tho 


r kernel 

1 «r itio 

Ik-an. 3S 

. Snmo 

early In 


lie lUl-'k .-..I 

tylflnni i 

Mng nn, 

1 KllMwl 

ng .lie iilu 

30, Snii. 

adcuK'ol in B<--rniiiia 

tloi.; Iho 

ok devel 

..l«.l in 

KXlo ..f 

■tern be 

ntlim n 




. 37. Embryo 



iiu UiG Meil-ciat. 

38. A 






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 cot3'ledons have 
not onl}' 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 
b}' the same organ, it rarel}' i)erforms l)oth 
equally well. Here the}* become mere 
receptacles of prepared food, the nature 
and office of which is the same as of the 
albumen, or nutritive dei)osit 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 
deix>sit. The plumule in such cases is 
alwa\'s apparent before germination ; and 
it develops even with more vigor than in 
the preceding cases. It usuall}' rises as a 
stout stem of several internodes lengthen- 
ing almost simultaneousl}', or at least the 
upiKjr strongl}' 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 slightl}' ; 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 hypogcBous (t. e. underground) situation of the cotyle- 
dons throughout the germination. The slight elongation of the 
caulicle senses merely to protnide its root-end from the coats of 
the seed in a downwanl direction, and from this a strong root 
usually is formed. 


FIG. 3D. SecUon of an acorn, ftlle;! by the embryo. 40. Ailvance^l germiimtlon of 
the tame. 

33. In some Oaks, notably in our Live Oak (Qnerens vire 
and loss so in tlit- llorstfbestuut, the two cot.vlmluns cwak-sLt or 
t^ohtre by llivir oontiguous faces. 
In some of these cases of liypo- 
gieoua germination, tlie sliurt 
caiiliule and pliunule are extri- 
cated IVom tlie enclosing coats or 
busk by the develo|)Uient uf short 
stalks {petioles, 157j to the fleshy 
cotyledons; asisseen tnFig. 43, 
and in most germinating auoms. 
These petioles are not visible iu 
tlie seed, but are the first develoi>- 
meut in germination. 

34. There arc some curious 
cases in which, while the eaulicle 
remains short and subterranean, 
tlie cotyledons are raised out of 
gi'oiuid in germiitatiun by the 
formation of far longer stalks 
(petioles) than those of the 
*i *' Ilorsechestmit. A singularly did- 

guiseil instance of this kind is seen in Megarrhiza, a genus of 
Cncurbitaceous plants of Calil'omia and Oregon, remurkuhlc for 
their huge root. The laige seed has vcrj- thick and fleshy 
ootykflons, and a very short and stmight canlicle. In germi- 
nation, the whole seed is elevated, seemingly in the manner of 
the bean, upon a stout stem. One waits tbr u lung time exiicct- 
ing to see the cotyledons throw off tlic bursting husk and expand. 
OT else to put forth the plumule fVom between their bases. But 
at length the plumule m^es its appearance from an uiifsijected 
place, coming separately out of the soil. Remoring this, the 
slate of things represented in Fig, 43 is presented, — that of 
the plumule seemingly originating fVom the base, instead of the 
ajiex, of an elongated caulicle ! But on examination of tbo cleft 
(Voni which this proceeds, by making a section of the stem aUivc 
(showing that it is hollow), and finally hy sei>aratiug the eotylo- 
dona and gently tearing apart the two short stulks by which tliey 
are unite<l to then- stem-like support, it is found that the latter may 
bo divided into two (as shown in Fig. 44). evendown to the cleft 
below. This explains the anomaly. The real cauHcle has ii;- 



mained short and subterranean, and is confluent with the upper 
part of the thickening root : the seeming caulicle, which raised 
the cotyledons 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 germinatidn 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 
compose<l : 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 cot3'le- 
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 
Cucurbitaceffi to 
bring up the cotyle- 
dons that they may 
develop as leaves 
(as in the Pumpkin, 
Fig. 47) : here this 

elevation is brought ^ « 

about in a different way, but without securing the useful end.^ 

» It may be inferred that Meganrhiza is a descendant of some Cucurbitacea 
with thinner cotyledons, which in germination developed into long-stalked 
leares, in the manner described in the next following paragraphs. 

FIO. 43, 44. Pecnliar germination of Megarrblza Callfornlca; explained above. 


3a. This same anomaly, as to the development of long stalka 
to the cotyledons and their union into a stem-like body, occurs in 
various stwcies of Larkspur (notably in the Catifomian Delphin- 
ium imdicaule) ; but in these the cotyledons develop into a pair 
of elBcient green leaves. 

3C. A similar elongation of petioles of the cotylwlons, but 
without any union, owiirs in u siwcies of Morning Glory of the 
plain.1 iK'yond the 3iIissisMippi (l|»oinffia leptopliylla) ; the leaf- 
like wityliilons coming np on their long stalks separately from 
the gntnnd (Fig. 45) ; the develo)>ed pbimule rising some 
time afterward between them. Com|)are this with the ordinary 
species (25, Fig. 15, 16, 25), and note that the difference is merely 
that the eaiilicle in the common Slorning Glorj' elongates and the 
I)etiiilcsof the cotyledons remain short. 

37. In all instances thus far a single primary root so rcgiiUirly 
devcloiis from the lower end of the axis of the embryo (vnriimsly 
named radicle. or eaiilicle), and forms snch a direct downward 


protongation of it, that it was called the dcscencting a^ ; and 
tbe body from which it originates was named tlie radicle, on the 
supposition that it was itself the nascent root. But, as alrcadj- 
cxplained, the ao-callnl radicle grows in tbe manner of stem (24), 
and is inoii)hologically that initial internoilc the node of which 
bears 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, Echinoeystis, and the like, 
Ihc strong cauliclc semis out directl}' from its root-end a cluster 
of roots or rootlets, of equal strength ; i. e., it stiikea root in 
nearly the manner that a cutting docs. (Fig. 47.) 

38. The Foljeotfledonons Embrjo is one having a whorl of 
more than two seed-leaves. The dicotyletlonous cmbiyo being 
a whorl of the very simplest kind, that is, with tlie 
members re<Iuced to two, the potyeotylcdonous 
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 ap[>car instcatl of 
two. In Knes (Kig. 48, 411) , however, ami i 
most but not all Conifera;, a whorl of from 3 to 10 
cotyledons is the normal stmeture, varying accord- 
ing to the species, but of almost uniform number . 
in each. In germination these are brought out of \ 
the soil bv tbe 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 line 
tribe, all the subsequent leaves are alternate (spiral) in arrange- 
ment, with some disguises. In the Cypress tribe, the cotyledons 
are fewer (not more timn four, and more commonly only two), 
and the subsequent leaves also are in whorls of two to four ; 
I. (., are either opposite or verticillntc. From the occasional 
union at base of the cotyledons of a polycotylwlonoua enibrio in 
pairs or groups, and from a study of their early development, 
Duchartre ' plausibly maintains that such cotyledons really consist 
of a single pair, partctl into divisions or lobes. The ordinary 
interpretation, however, is equally tenable. 

39. The Monocfltfledonoos Embrjo, although theoretically the 
simplest, is practically a more difflciilt study. It has a single 
cotjledon (as the name denotes) ; also a single leaf to each node 

' Ann. Sd Nat. ser. 3, x. 207. This view, which originated wUliJuasieu, 
b «ilopl«d by Parlalorc in DC. IVodr. xvi. 

FIO. «. S«etl™ofinw«lot»Plne.wlllill«rnilirvniifii.'verslcotyIeaoM, 48. Early 
MHlUntt P1n«, trilli lu •tamlel, dliplayinn in uli wwl-luat™. 



of the plumule ; that is, the leaies of the embrj'o are alt«mS 
But the oauliele is usually icrj- short, and there is no estcrtial 
mark by which its limits inaj- be ilistiii- 
Buishe<! fVom the cotyledon, until genni- 
jifttioii has l>Jgmi, For a tj-pc of it. the 
cnibrj'o of some aquatic or marsh plants 
I may lie taken, whore it forms thi? whole 
I kernel of the seed (Fig. 50-53), and 
the structure can he made out autecvdeiit 
to germination. It is understood by Bupjiosing that the cotyle- 
don, whieli forms its prindpal bulk (the caulJele being only tlio 
very short thickish base) , is couvulute around a short 
I plumule, and the margins concreteil, except a minute 
I longitudinal chink at base, out of which the growing 
])lumulc protrudes in germination. The embryo of 
I may be similar in structure, but no distinction 
of parts is visible. It is very small in proiwrtion to 
the size of the seed, the kernel being mostly albu- 
men. — a supply of footl, tVom which the genoiuating 
cmbrj'o 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 fonns at the fVee end of the exeesMvcly 
short caulicle, and the plumule derelops from the 
other in a series of one-leaved nodes, the intcniodes 
-^-ji of which remain so short that the leaves continue 
'^/jf in close contact, the bases of the older sueceesively 
Ij enclosing the inner and younger. (Fig. 56.) Here, 

therefore, Uie cotyledon mainly remains in the seed, 
and the seed remains underground (hyiwgtcous) . 

40. It is somewhat different in the Onion, which 
has a similar embryo, except that it is louger, and 
the cotyledon is curved in the albumen of the seed. 
The lirat steps arc the same as in Iris ; but as soon 
as a root is formed and embedded in the soil, tho 
cotyledon lengthens vastly more, into a long and 
filiform green leaf, which, taking an erect ijosition, 

UxA at Trleloehln puliuln ; tlie rhaptic. Isntlng to Ibe ilruiie cliatuaM tha 

e<1 towitnU tliD e7« Al. Tlio rnbryo iletuhAl fn>mlhuBeml.coiit«, «hi>wln( 

rt!i8«nylorton: llionhortiarl below 1*lliet»i]|p]e. 

orally, arut half Itin coIxIrI^ cm awar. brinKlng to 

ooUTMlffll wfUiln. 03, A crota-iBPUDii thraugti tbo jilumulp, mora 


carrica up the light eeed far above the surface of the ground, the 
tip only remauimg in the albumen of the seed until that is ex- 
hausted, when the tip perishes and the emptied husk falls away. 
Ahout thia time the plumule shoots forth from one side of the 
subterranean base of this t-otyledonar leaf in the form of a second 
and simdar filiform leaf to be followed b\ a third and so on. 
The sheathing bases of these suocee<lmg lea\es become the coats 
of the OoioD bulb The mtemodes remain unde\eloiKKl until the 
plant IB read} to blossom Veiy similar is the gci-mination of 
a date-seed except that the 
protruding cotj ledon does 
not lengthen so much nor 
does it eleiate the hea>\ I 
seed Instead of the seed i 
being earned up the lower 

end of the embrj o contain " » w 

ing the plumule is pushed down more or less into the loose 
soil from which m time the developing plumule emerges. 

41 The cmbno of Grasses cspeeiall} of those which yield 
the cereal grains is more complex owing mainh to the great de- 
velopment of the plumule 
and the manner in which • 
its nidimentarj leaves 
Buccessuelj enclose each 
otbor That of Maize or 
Indian Com one of the 
largest, 18 most convement 
for study. (Fig. 56-59.) The floury part of the seed, which 
makes most of its bulk, is the albumen, lai^ely composed of 
starch. The embrjo 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 com here represented. Tlie 
form of the embrjo b best sbown, detached entire, in Fig, 58 ; 
its structure appears in the sections. The outer [rnrt is the 
cotyledon, which incompletely enwraps the i)Uimule : it adheres 
closely to the albumen by the whole back, and remains un- 
changed in germination : its function is to absorb nutritive 

FIO. EC Section, flHtwlH, of ft grain of InilJan Corn. iliiMIng the Blliumcnuid tlie 
mbryo. or. Similar iHtlun at rlghi luinin lo Die tint, H. A ileUclied enilirfu^ 
oormponillni puU ot Pig. SI ami M Inillcateil by ilDltal lino. 

Fia. B9. Vartlcal Kcllon of Inllan Com un»> I1>e ilikkiicxs of Die gnln, Jlvliting 
tlie embryo through tbe centre ami dlnpLayhig lie partj: c, ootylvdon; p plumuloi 
r, tlie mild* or canllcle. 

Pia. 00. Similar lecUan of gtnia of rice. 61. Same of an oat^gralni tbe parti 


matter rumished by the nllmnieu, and to tronsmit it to t 
growing i)luiniile. The ]iluiuule conaista of a succession of 
ru<liinen(srj' loave§. slieatliing 
and L'liL-lusing ouu auolhur. on 
the stinuuit of a ver^- short 
axis, whicli is mainly the 
oauliolo. otliorwiao fnlltii niil- 
icle. This is ciimplotvly en- 
closed by a basal (rartioa of 
the cotyledon and of the 
outermost leaf of the plu- 
mule, which Torm a peculiar 
sheath for it, named tlie 
Cu/forAira,' i, e, root-sheoth : 
conseqneutly the first root or 
i-oots have to break through 
this covering. As in Ilie Oak 
and !*eB(a2). the very lirst or 
" outennost leaves of the plu- 

mule develop impeifeetly and not into 
ellicient foliage. The one in Kig. C2, 
which encloses the rest in the early 
growth, is left bi'hind as a mere sheath 
> to thf base of the following and more 
perfect leaves: it is the same as the 
lowest in Fig. C3, Tlic leaves arc first 
developeil : the intemodes lengthen later, and the lowest lengthen 
very little. Xot rarely Uie first root starts singly from the tip of 
Uie caulicle (Fig, (i2. Just as in Fig. 55) ; but others of equal 
strength follow from any jtart of the cauliele, and soon iVfim 
the nodes above ; and no tap-root is ever formei). 

42. A Pteuilo-monocoltilfdonout erahrjo oceHsioually occurs ; 
that is, one of the dkiityledonons ty|K>. of wliieli one cotyledon is 
wanting through aliortlon. This occurs in Abronia, a genus 
related to Mirabilis. and bearing an embryo veiy similar to that 
representt-d in I'ig. I 7. I'*, e.toept that one cotyledon is ul«ent. 
The anomaly of an acoiyledonuiu embr\o oecurs in Dodder, a 
plant of the dicotyleilonous typo, but with botti eotjleduus 

> ThU. tlic Colrothiif of MirlM>l, iliuuld not be cnnrnumU'il (>» by *otiui it 
hu be«n) witli Ibc "nxit^ap," nr tiBSuc vflikh onllnnry mutB (wliftlii-r 
primary or *(vuuit>ry) brvwk tliruiigli in iheir dt^Tt^lopnivnl or carry on 
iheir apex. 

FIO. CS. Entlir lernilnmlnn <if Imllwi Cnni. flS. Mi>re vlrancol nniiliintiuii et 


actually wanting, — a correlation with ito parasitic mode of life. 
(64, Fig. 78.) 

43. The dicotyledonous and the monoc-otylL><lonous character 
of the embryo is correlated with profound differeucc» in the whole 
ulterior development, as revealed in the striietnrc of the stem, 
leaves, and flower ; which differences mark the two great divisions 
of Phxnogamous plants, viz. DicorvLF.nosES or DicoxvLEWoxoiia 
Plast3, and 6Iosocotyledoses or Moxocotiledonous Plants, 
— names introduced into classiticatiou by Ray, aud ado[)ted by 
A. L. Jus^eu, in his Genera Plautarum. 



Section I. Of tue Rdor. 

44. The Root, which has been called the descending avis is 
that portion of the bo<ly of the plant whith gions downward, 
ordinarilj' fixing the vegetable to tliL soil and abboibing from it 
materials which the plant may elaltoratc into nounshnKut Aa 
already stated (24), the root gmws m length b^ continuous 
additions of new fabric to its lowei e\tn,mit\ elongating from 
that part only or chiefly ; so that tlit tip of _ 

a growing root always consists of tin nio^t 1 

newly formed and active tissue. It minmlh ^ 

begins, in germination, at the root tnil iil tin, 
caulicle, or so ealletl radicle. But rooti so m 
proceed, or may proceed, from otlicr |)arts of 
the stem, when this is favorably situiitiil f >i 
their production. The root docs not grow ^ 

from ita naked ayicx, but from a stratum ' ^ 

immediately behind it: conscqiKtith its blunt at' 
or obtusely conical advancing tip <onsi8t3 of oldtr, finner, and 
in part effete tissue. The tip of all secondary roots and i-ootleta 

FIO. «. MagnlOed tip nf root of m Kolling Maplo (»ucli u in FJg. Dl. .ufflilcntrjr 
mUried to InrtlcaU the cellular ■tniclarc: a. Iha poitlon nlicn srawUi i) taking 
plB»i b. the older uul Onnor ll|i. 



ii similarlj- capped or protected.' But the Bo-called root-tap ia 
seldom so distinct or eojuirable as to dL>sen'e a [)articular name. 
45. Katnre of Qrofrtb, Cells. The (levclopmcnt and growth of 
tha root, aa of other oi^aiis, results from the development, 
growth, and increase in number of certain miuut« parts, of which 
tlic plant ia built up. These component parts aro so much alike, 
at least in an early stage, and aro so obiioiisly forme«l all on 
one typs, that tlicy take one commoo name, that of Celx£. 
ThiBO aro the histological elements of plants, i. e. the units of 
minute anatomical structure. While, in the mori)hol<^- of the 
plant's obvious oi^ans, analysts brings us to the phyfomer (IG) 
as the individual element which by a kind of propagation 
jtroduces its like in a second jihytomer, remaining however in 
connection with the first, thus building up the general strueture, 
80, in an analogous way, each of the obrious parts — each stalk 
or blade or rootlet — is microscopically detormiiicd to be com- 
posed of these ultimate oi^anic units, generally called cells. 
The ceU {crUula, by the French 
conveniently termed cellule) is the 
living vegetable tuiit, in the same 
sense that the brick is the unit 
of a brick edifice. To make this 
analog}' fairly complete, the 
;k3 should be imagined to 
have a fiim exterior or shell, 
and a soft or at length hollow 
- interior, also to be living when 
incoiiKH-ated into the structure, 
an<l finally to he [troduecd in the 
\ forming stnicture by a kind of 
propagation. The ])ro<luction or 
increase in nnmber of these cells by development from previous 
on:>9, an<i their successive increase in size up to maturity, are 
what constitutes vegetable grounh.* The in8]%ction through a 

■ Tlic notion (tiat Ehc lip of the rool conaials of ilelioatc forming or 
newlj tormccl tigsur. or bean lomc organ or itructurv of lliis nature (a 
"Ajj«ni7iWc"),liafi hanllyyi.'llH.-oni'1iinina(c<]from llic li.'xt.|>uuk» ani] popular 
writing*. It ha<] no propiT founilation in fact. 

In I^mmt, and In aomu oilier nqualkf, anil aUo in somp aerial roote. thii 
oUlcr liuue often icpartttca into a real root-cap. tree at bote, like an inrorted 

' Tliw. ai to Ihc alructuro, in the aubJMt of Ilitlnlirji/ : na lo proccsaes or 
actiona, the aubjrct of Phi/tiiJirig ; both lo be (reateO in a acparalc volume. 

no. U. M, Pnrtlnna nf aurfore at FtR. 04. innre mtsnlflni. clfarly ill*p1aylnK Ihs 
■npurBi-inl rrlliilur BtrnPiure ami ihe InnK pnv^wa ftom ■onii! or tbg c«lla, calleJ not- 
Ajin, wlili'li abounil on Uis iip|nr t«rt oT Fig SL 


simple microscope of a slender 3'oung root, and of thin slices 
of it immersed in water, may serve to give a general though 
crude idea of the vegetable cellular structure, sufficient for the 
present purpose. Roots are naked ; that is, they 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 predetennincd oixler. The ultimate and 
very slender branches are sometimes called root-fibrils ; but 
these are onl}' 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 readil}' 
when 3'oung and tender, less so when older and finner. 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 b}' the spread of foliage, so in a less 
degree is the absorbing surface of young roots increased b}* the 
production of root-hairs. (Fig. 64, upixjr part, and more magni- 
fied in Fig. 65, 66.) These are attenuated outgrowths of some 
part of the superficial cells into capillan' tubes (only one from 
each cell) , closed at the tip, but the calibre at base continuous 
with the cavit}' 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. 
P^urther back the older or effete root-hairs die awa}' 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 leafy 
branches. Although not naturally- furnished with buds in the 
manner of the stem, yet many roots have the ix)wer 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 are wounded. 
And the roots of Madura or Osage Orange so readily originate 
buds that the tree is commonly propagated b}' root-cuttings. 

48. Kinds of Roots. The root, commonly single, which origi- 
nates from the embryo itself, is called the PmMARv Root. (37.) 
Roots which originate from other and later parts of the stem, 
or elsewhere, are distinguished as Secondary Roots. But the 


latter are as normal as the primary root; that is, to stems 
so situated that they can procUice them. Most creeping plants 
emit them freel}*, usuall}' from the nodes ; and so do most 
branches, not too old, when bent to the ground and covered 
with earth, thus securing the rcquisite moisture and darkness. 
Separate pieces of 3'oung stems (cuttings) can commonh- be 
made to strike root. Ujwn this faculty of stems to originate 
roots depends all propagation by division, In' lading or la^'ering, 
by cuttings, &c. It is mainly annuals and common ti*ees that 
natural!}' dei)end on the primary ix)ot ; and most of these can be 
made to produce secondarv 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 ix?si>ect to this, either the 
root or the plant, as the case may be, is distinguished into 
Annual^ Biennial^ or Perennial, acconling to whether life is contin- 
ued for a single 3'ear or season, for two, or for a greater nmnbcr. 
The diffeixjnce is not in all cases absolute or even well marked. 

50. Annuals are plants which, springing from the seed, flower 
and seed the same vear or season, and die at or before its close. 
The}' produce ^^0M« roots, cither directly from the embr3'o and 
succeeding joints of stem (as in Grasses, Fig. 03), or from a 
IMjrsistent primary' or tap-root^ more or less thickened into a tnmk 
or divideil into branches. The products of vegetation in all such 
herbs are not stored in suliterrancan or other reservoirs, but are 
exi)ended directly in new vegetative growth, in the production 
of blossom, and finallv 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 existonco prolonged b}' 
not allowing them to blossom or seed. Othei-s, with prostrate 
stem or branches, may from these produce secondar}* roots, 
which, fomiing new connections with the soil, enable the newer 
growth to sunive when the older parts with the original root 
have perished. And many herbs, naturally annuals, are continued 
from year to year through such propagation from the branches, 
use<l as layers or cuttings. IMoreover, certain plants (such as 
Ricinus or Castor-oil Plant), which are 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 chmates with comparatively warm and rainy 
winter and rainless summer, many germinate in autumn, vegetate 



through the winter, flower and seed in spring, and perish in 
eariy summer. These may he termed Winter Annuals. 

53. Blennlala are jjlants which, springing Trom the seeil and 
^-egetating in one season, live through the interruption of winter, 
and blossom, fnictifi, and i»erish in the next growing season ; 
tlieir lire being thus divided into two stages, the Rrst of vegeta- 
tion, the second of fnictiflcatioii. In typical biennials, nearly 
the whole work of vi^tation ia 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 verj' short stem in e juneetion with tlie root. The 
root of a biennial accordingly enlaiges anil becomes fleshj", or 
obeB3, as this matt«r accumulates. At the close of the growing 
season, — the leaves perisliing and the 
stem having remainetl verj- short (with 
undeveloped int^modes), — the root, 
crowned witli the bud or buds, contains 
the main result of the summer's work, - 
aa provision for the nest j'car's devel- -,_> 
opment and lh2 completion of the 
ej'cle. This development, being thus 
amply provided for, is undertaken in 
spring with great vigor ; blossom, fruit, 
and see<l are rapidly produced ; and 
the stock being consumwl, but not at 
all replenished, the cells of the great 
root arc now empty and effete, and 
the individual perishes. The Beet, 
Turnip, I'arsnip, and Carrot are fa- 
miliar examples of biennials, with the 
store of nourishment in the root.' " 

The Kolil-ral)i is a biennial with this deposit in the stem : 
the Cabbage, iwrtly in the stem, partly in the head of leaves. 

> In these the cauliclu cnUrgos with the roal, so that the uppLT and 
bud-braring end i> Btcm. 

Tap^wU of thij kind «re laid, in deseriplive botany, lo be 

Fiai/nriH or Spintltt-ihnpril, when broiider in the middle atid tapering 
towarils bold endi, as in the common Kudiali [i'ig. ITi); 

Conical, when taperiniT reRUlariy from base lo tip. as in eairots. &c. ; 

Kiipi/orm, i. e. Tumii>-ihiiiie'l,vhen the thickentil part is ividtr tlian liijih, &c. 

Fiudrfrd RoMi are Ibose which form in cluslora; tliosc may be slendtT or 
thickened. When much tliickoned, either irregularly or not of the abovu 
•hapei, tlicy are taid to be tuUmut. 

no. «I BailJsli: a rii«ir.rm ia|>-rwl. 


54. BdI some plaoU. such as tbe Rsdish, whicb when t 
spring from seeil iu autumn are trae biennials, will when raim-)l 
in spring pass on directly to the IloweriDg stage in suTnincr. c 
when sown after the warm season Ik^qs will often nia ihrough 
their course as annuals. Then there arc varions biennials whieh 
thicken the root very little and hold their learoa through the 
winter. Between these and winter annuals no I'leur ilemarcatiou 
pan be drawn. As rcsijoetd annual and biennial duration, (he 
terms may for the most part be applied indiscriminfllcly to the 
plant or to the root. We nmy 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 percnuial 

roots. In trees and sliruba. also in 
lieibs with growth from year to j-car 
from a strong tap-root, the root 
is naturally perennial. But in most , 
perennials with only Sbrous roots, 
these are produced anew tVom timo ' 
to time or from year to year. Also, 
^ while some such roots remain flbrous 
nil sene only for absoqitiou, others 
\ may thicken in the manner of the 
, ordinarj- biennial root and serve a 
r similar nse, i. e. beeome reservoirs of 
elaborated noitrishmeut. The Dahlia 
(Fig. (18) and the Peony afl'ord good 
examples of this. iSweet potato ia 
another instance.' Klost such roola I 
have only a biennial duration : tbcy 
are proclnced in one growing season; they yield their store to 
fonn or aid the growth of the next. When perennials store tip ' 
nutritive matU-r undergrouml. the deposit is more commonly 
made in a subterranean portion of the stem, in tubers, eornu, 
bulbs, &o. (See 115-122.) 

50. The distinction between annuals and biennials is at timc« 
80 difUcult. and the particular in which they agree so manifest, 
— namely. Hint of blossoming only once, then dying, as it wcrp 
by exliaustion, — that it was projwsed by DcCandoUe to unite 

1 It la na\y hy the reaillnr** o( lhi« ront to produce arlvtriltlioiis buit*, 
cipeoidlly (mm itt upper ixti, tliat U h«» lieen minlakcn fur a tulicr, »ui;li 
M the cummuii polnto, 

FtU. W. Pu'lcli!dan(ltutcrouiorrUiiirotm(iMandai7}r«>l*oriMiU>i i 


the two ander the common appellation of Monocarpic plants, 
PlarUa monocarp%c€B^ taken in the sense of only once-fniiting 
plants ; and to designate perennials by the corresponding tenn 
of PoLYCARPic, Planttp polycarpica^ literally many-fruited, taken 
in the sense of man3'-time8 fruiting.^ 

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 six?cies 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 sur\ive the winter ; but whenever fnictified 
the fibrous-rooted mother plant dies throughout ; yet usually not 
before it has established, and perhaps detached from the l)ase, 
small offsets to blossom the next season ; and so on. Then 
Houseleeks (Sempervivum) and such-like fibrous-rooted succu- 
lent plants multiply freely b}' offsets which are truly perennial 
in the sense that they live and grow for a few or several 3ears ; 
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 sunnving. 
The same is true of the Centurj' plant (Agave Americana, 
wronglj- 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 usuall}' sends forth roots only when 
covered by or resting on the soil, which affoixls congenial dark- 
ness and moisture, yet these are in some cases produced in the 
open air. Roots may likewise subser\'e other and more special 
uses than the . absorption of crude or the storing of elaborated 

59. Aerial Roots is a general name for those which are pro- 
duced in the open air. One class of these may ser^•e the ofllce 
of ordinary roots, by descending to the ground and l>ecoming 
established in the soil. This occurs, on a small scale, in the 
stems of Indian Com ; the lower nodes emitting roots which 
grow to the length of several inches Injfore they reach the ground 

* 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. Mondforous (bearing progeny once) and Polytocous (bearing 
many times) would be more appropriate. 




into which they i>enctratc. More rcmarknble cases &bodm 
those tropical regions where the sultry air, eatumted wiih moist- 
ure for a lai'ge jiart of Uic jear, favoi-a the utmost luxuriance of 
vegetation. In the Pahn-Uke Paudanus or Screw-Pine' {Fig, 
CO), verj- strong roots, emitted in the open air IVom 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 
(Vom the horizontal 
branches of the tree, 
often at a great 
height, atfiretswing- i 
ing IVee in the air, 
bnt Anally reach- 
ing and establishing 
themselves in the 
grouud, where they 
increase in diameter 
and form acceasory i 
tninka, surrounding 
the original bole and snpixirting the wide-spread canopy of 
branches and foliagu. Verj- similar is the economy of the Man- 
grove (Fig. 71), which forms iinijenetrable thickets on low and 
muddy sea-shores in the tropica throughout most parts of the 
world, extending even to the coast of Florida and Louisiana. 
Here aerial roots spring not only fhsm the main tnink, as in 
the Pandnnus, but also from the branchlets, as in the Banjan. 
Even the radicle of the cmbrjo starts into growth, protrudes, 
and attains considerable length while the fruit is atill attached to 
the branch. 

50*. Aerial Rootlets for pllmbln? are familiar in the Ivj~ of the 
Old World (Hedera), Tniin|iet-t'reeper (Tecoma radicans), and 
our Poison Ivy (Rhus Toxicodendron) ; by tlie adhesion of 

' So nnmcil, nol from «ny rcictnblBni^c to « Pine-tree, bat from m like- 
neBi at the foliage la that of ■ Pine-Apple. 

FIO, en. PuniUiiu*, or Bcreir.pluoi utd lii U» twikgraBni), TO, ■ 

tKli<io|.lioni Muiglr), 




60. EpIiArtM or Air-Plants also hnvp roots which aro tlirongh- 
I oiit life unconnected with the ground. Epiphytei. or Epiphyiie 
[ plants, as the name dentjtx's. are such a« grow upon other plants 
T mthatit taking nourishment IVora them. Deriving tliis from the 
I air nlonc, they arc culleil Air-jAanU. Tliie name might be 
I extended to thv &.im(> or other ktnda of jilants attaebiiig them- 
I selves to hare wbUs, rocks, and the like, and un(.-onneotG<l with 
I Ul« soil, tliuugli sncli would not tcehiiically be epiph,^*tl.■s. Very 
I nan}- Lichens, Mosses, anil other plants of the lower grade, and 
r not a few phienogatnoiis plants, are in tliis case. The greater 
I part of Uie phicnt^atnous Epipli^'tea |>er1ain U> two munocotyle* 
I lJunoiisonh'rs, the Orchis family and that to which the Pinc- 
I Apple liolongs, viz. the Bromeliaceie. Their thread-like or 
I cord-like simple roots either adhei-e to the baik of the supporting 
P tnw, secnrini; tlw plant in its ])ositiDn, or siitn« Iiniig loose in the 
Of thrsc. Orchids, 1. 1. plants of tlic Orchis family, are the 
I most showy and numerous, and of the greatest variety of forms, 
I «i>]K.-nBll,v of tlunr blossoms, which ar« oHen bizarre and fantjis- 
Thcy >>elong, natnriilly, to climates wluch are hotli warm 
I and bimiid : they are highly prized in hot-honse cultivation ; 
I lunl, aloi]g wilii tlie bnrily and terrestrial [lortion of tlio order, 
I they are peculiarly interesting to the botanist on account of the 
I cuigular and exqnisite adaptation of tlieir flowers in i-elalion 
(luiDiir«tii which visit them. In some the blossoms cunonsly 

FIO. II TUo Bud 



resemble butterflies or other insects ; as, for extunple. Onciditnn I 
PapUio, Fig. 72. Epiphytic orchids are iDtllgcnous to the Unit«tl 
States only from Georgia to Texiis, and oaly in humble fomis, 
iu company with stx^cies of Tillandsia. representing Bromeliacc- 
ons cpijihytca. The commonest of the latter tribe, and of most 
nortliem range, is the T. asneoidcs, the so-called Ivong Moss, 
which, pendent in long and tangled gray clusters or festoons from 
the branches of the Live-Oak or Long-leaved I*ine. gives such a 
peculiar and sombre aspect to the forests of the warmer portions 
of our Southern States. ^ 

CI, ParAsillc Plants have the peculiurity that their roots, or i 
wlmt answer to roots, not only fix tliemselves to other plants, 
but draw Ihurefrom their noiuishment, at least in purl. Among j 
crji>ti:^nmou3 plants verj' many Fungi arc parasitic upon or I 
within living plants or animals. But only phienc^amous para- [ 
sites are here under conaidemtion. These may be divided into J 
two classes ; those with and those without green foliage. 

G2, Qrwn ParatiltM may be either wholly or partially parasitic j 
that is. thi'V mny draw all tlieir supjiort from a foster |>lant, or 

, Iwg lIloWJIGpllllll^MOrilie 



thej- may be likewise rooted ia the soil, and receive from it 
materiab of their food. Having green foliage, they are capable 
of elaborating such food, whether taken directly from the soil or 
from the crude sap of the foster plant. The Mistletoes (Viscum 
and its allies) are the principal examplea of complete green 
parasitic plants. Seeds dropped by birds on the boughs of trees 
germinate there ; the root-end of the caulicle jwints thither instead 
of towards the earth ; the root, or what would be such, pene- 
trates the bark and in- 
corporates itself with the 
aap-wood so perfectly 
that the junction of par- 
asite with foster trunk 
is like that of branch 
with parent trunk. The 
parasite is probably fed 
by both elaborated and 
crude sap, that ia, both 
by what the foster tree ^ 
has assimilated and ^^ 
what it has merely taken 
from the soil and air: 
the former it can at once , 
incorporate; the latter ' 
it has first to assimilate ' 
in its own green leaves. 
Sometimes one Mistleto 
is parasitic upon an< 
other of the same or of 
a different species. 

63. Partiallypara8iticplants(mostlygreen) maybe either woody 
and arborescent or herbaceous. The species of Clusia in tropical 

FIO, 7«. NsUtc eplplijtel of GencKla. £1; : tli« ertKt oiia nt tliB rlglil 
EptdoDdnuB smHpmiD; tbe banglni nne Tlllaniliila aiineiJiln. cMti\ Lnn 

Fia. TIL Boota of a«nrdl* fla»: ■nms of tlis nxtlvts ■ttaclilnfthomi 
Mlcall; totluniiittfaBlnsbciiTy. (From ailrawlng bj Mr. J. Stauflbr.) 


America (called Cursed Fig) are examples of the former. They 
form trees, send down aerial roots in the manner of tlie Banyan ; 
but, wliile some roota seeit the giwimcl, aome may attach tJiein- 
aelves to other trees parasitically, and draw from them a portion 
of their support. The iiarositism of certain herbaceous pknts 
with green foliage is clandestine, the conuecUon being under- 
ground and therefore long unsus|)ected. This occurs in species 
of Gerardia (at least of the sectioti 
Dasystoma) and other plants of the 
same family, the uncultivability of 
nhich is thereby explained. Also 
in Comandra and in their relatives 
' Ihe Thesiums of the Old World, 
belonging to a iiatnral order (the Santalaceie) which has much 
affinity with the entirely parasitic order (Loranthacea;) to whiuh 
the Mistleto belongs. 

G4. Palo or Colored Farasitm, such as Beech-Drops, Fine-Sap, 
&c.. are those which are destitute of green herbage, and are 
usually of a white, tawny, or reddish 
hue ; in fact, of any color except 
green. These strike their roots, or 
sucker-shaped discs, into the bark, 
mostly that of the root, of other 
plants, and thence draw their food 
I'rom the sap already elaborated. 
They have acconlingly no occasion 
for digestive organs of tliuir own, 
t. e. for green foliage. The Dodder 
(Fig. 77) is a common plant of 
tins kind which is parasitic above 
ground. Its seeds germinate in 
the eailh. but form no proper root : 
when the slender twining stem 
reaches the surrounding herbage, 
it forms suckers, which attach 
themselves finnly to the surface of the supporting plant, 
penetrate its epidermis, and feed upon its juices ; while the 
original root and base of the stem [x^rish, and the plant has 
no longer anj- connection with the soil. Thus stealing its nour- 

FIO. T6. SHilnnofaneDniie atticheil molleUoT OurHnlla. ihovInK tin unlnn. 

PIQ. TT. TtiE mmniun DniMet of Ibo NorllNrn StaUa ICiucuU OrunuiUl, uf tlia 
nMunI tlia. jATwllIc upon Uie uea of ui bsrb: Ihe uiuwUihI gnrllnn mttba Inwn end 
nhawt tlM tDiHlE uf lu ■itaebmcnt. 11. Tbo call«d unbryo tikta from ilia tat. con- 
■l^iigofntkoit cftullcleuid (ilntsnle; moilemuty macnlflBcl. TS. Ths nine in edtdiI- 


ishmeat ready prepared, it requires no proper digestive organs 
of its own, and, consequently, does not produce leaves. This 
economy is foreshadoved in the embrj-o of the Dodder, which is 
a naked thread spirally coiled in the seed (Fig. 78, 79), and 
presenting no vestige of cotyledons or seed-leaves. A species 
of Dodder infests and greatly injures flax in Europe, and some- 
times makes its appearance in our own flas-fields, haWng been 
introduced with the imported seed. Such parasites do not live 
upon all plants indiscriminately, but only upon those whose 
elaborate Juices furnish a propitious nourishment.' Some of 
them are restricted, or nearly so, to a particular species ; others 
show little preference, or are found indilferently upon several 
species of different families. Their seeds, in some cases, it is 
said, will germinate only when in contact with the stem or root 
of the species upon which they are destined to live. Having no 
need of herbage, such plants may be reduced to a stalk bearing 
a single flower or a cluster of flowers, or even to a single blossom 
developed from a bud directly parasitic on the bark of the foster 
plant. Of this kind are the several species of I^lostylea (para- 
sitic flowers on the shoots of Leguminous plants) in Tropical 
America, one species of which was discovered by Dr. Thurber 

De&r the southern borders of New Mexico. Its flowers are 
small, only about a quarter of an inch in diameter. The most 
wonderfld pUnt of this kind ia that vegetable Titan, the Raf- 
flesia Amoldi of Sumatra (Fig. 80) which grows upon the stem 
of a kind of a Cissns or Vitis, It is a parasitic flower, measuring 
nine feet in cireumference, and weighing fineen pounds ! Its 
color is light orange, mottled with yellowish-white. 

> UoDotnipa or Indian Pipe (anil pcrhapi snme ri'latetl planls), allhough 
prab«bl7 pBTuilic oBliTin((rootiinc»rly gTO«lli,appt»ri lo live alterwanls 
in the maiiDer nf the larger Fungi, upon leaf-moulU and decaying herbage, 
lu mode of life ibonld be inveiligited. 

a bsd, cllrectlr parulllc on 


Sectiun II. Of Buua. 

65. Buds are the genns of sterna : they are asos with tlieir 
appciKloges in an tuilj stale, Leaf-bcds (Geuu^.) are those 
devolMl to vegetation, mid the |)aite, or some of tiiem, develo[) 
as leaves. Mixed bids eoiitaiu both foliage ami flower or 
flowers. Floweu-buus (ALAbA&TitA) are unexpanded blossoms. 
These are eoiisiilered in another ehnpler. 

66. The conspicuous ]>orttoD of auonliiiaiy bud. or that whieh 
first develops, usually consists of leaves, or scales tlie huraologues 
of leaves; the axis itself being very short and uudevelopcd. If 
this remains coin|uiratively short, the leaves as develojwd are 
ci'owded in a rosette, as in a Houseleek (Fig, 91*), a Barberry 
and the Larch; when the iutemotlee lengthen, the leaves are 
interspaced upon the axis. 

67. The cotyledons and plumule of the embryo are, morpho- 
logic-ally, tlie flrat bud, on the summit of the initial stem, tlie 
cauliele. This in germination and subsequent gi-owth develops 
into a leafy stem, in the manner already deseribeil. Normally 
this stem has the cajtaeity of growing on in this way from the 
apex or growing point, which is always potentially a bud, the 
apical or terminal bud (15). l^^omctimes it is merely potential. 
aud there is no external structure visible until the new growth 
be^ns, or ttie bud is said to be latent. 

08. Butcommonly.inplants that live IVom year to year, growth 
is divided into seasons or stages, with inter\'als of repose. Id 
such eases, espeeiallj' in trees aud shrubs, instead of a continuous 
succession of foliage, the period of interruption is apt to be 
marked bj" the production of scales {Bad-scales, Perulm, etc.) or 
dry teguments, whieh serve to protect the tender rudiments or 
growing poiut within during the season of rest. This being the 
winter-season in cold climates, Linn^us gave to such bud-cover- 
ings the common name of HiBEHSAcuLfM. From tlie usually 
tqnamou (seale-like) eharaeter of this covering, such buds take 
the name of 

69. Scaly Buds. Large and strong ones of this kind, such as 
those of lloraechestnut. Magnolia, Hickory, Lilac, &c., may be 
taken as the type of bud. The sealcs scne to protect the ten- 
der parts within against injury from nioistnrc and from sudden 
changes in temperature during the dormant or earliest growing 
state. To ward ofi* moisture more elfectually, (hey arc sometimes 
coateil with a waxy, resinous, or balsamic exudation, as is con- 
spicuous on the scales of the Horsechcstnnt. Balsam-Poplar or 
Balm of Gilead, and Balsam-Fir. To guani against sudden 



changes of temperature, they are oftea lined, or the rudimentary 
leaves within invested with non-conducting down or wool. 

70. Kitare ot Bod-scales. That they answer to leaves is made 
manifest by a coneideratioD of their 
situation and arrangement, which are 
the same as of the proper leaves of 
the species ; and by the gradual transi- 
tions from the former to the latter in 
many plants. In the Turioag, or sub- , 
terranean budding shoots of aumcrous // 
perennial herbs, and in the unfolding / 
buds of the Lilac and Sweet Buckeye ■ \ 
(^sculus pan-iflora), every gradation ^ ,1 
may be traced tietween bud-scales and . 
foliage, showing that do line of distinc 
tion can be drawn between them, but ' 
that the two are essentially of the sa: 
nature, are different modifications of 
the same organ. In the Lilac they 
may be r^arded as the blade of the 
leaf, modified and depauperate ; in the 
Buckeye (Fig. 233), and therefore in 
Horsechestnutjas the base of leaf-stalks ; 
in Magnolia (Fig. 81, 82), in the 
Tulip-tree, and in the Beech, they are 
endently stipules. Tliey must therefore 
be referred to in the section on the 
morpliologj' of leaves. (227.) 

71. Naked Buds, tc, of shrubs and 
trees, even in climates with severe 
winter, are not unknown, that is, buds 
unprotected by special scales or other 
coverings. For example, the latest 
pair of loaves of the season in Viburnum 
nudum, V. lantanoides (Hobblebuah), 
and the like, remain in a nascent state 
over winter without covering, and ex- 
pand into the first foliage in the spring. 
Yet V. OpuluB (Snowball, &c.), another species of the same 
genus and inhabiting the same region, lias wcU-fonned scaly 

FIO. 81. Bnncb or HrkgnnlU Unbn] 
teimtiial biHl; ud below eihlblllnK >he li 
or luiDiilu ■nn Ml by Ihi fall of the ba>l-« 
ftcmie from a ilinilAT bufi ; Id 1hlcliene<t Mxl: 

le bue of * lMf-«tBUi ; tba m 



teof-buda. In other bartly shrubs and trees, the buds, equally 
or almost destitute of scales, 
are minut«, liutdvn in or un- 
der the bark, or othenriae 
incon8t>i(iious until vernal 
growth commences. Phibi- 
delpliuB aud Tuxoilium arc of 
this kind. 

72. Hnbpetlolar Buds. Some 
leaf-buds ai-e singularly cov- 
ered in their eaily slate and 
through tile summer, as iu tbe 
Locust (Robinia), Honey-Lo- 
cust Fig. 96 (where they re- 
in Vfry uiidevelojied) , in 
Yellow Wood (Cladrastisj.and 
I more conspicuously in the 
' Plane-tree (I'latanus, Fig. 87); 
here they are all formed un- 
der the base of the protecting 
leaf-stalk, which in Plane-tree 
forms asheathorinvertedcup, 
verj' likea eandle-extinguisher, 

fitted to and concealing the conical bud until autunm, when by 

the fell of the leaves these buds are exjiosed. 

no. S3. tMnsnin nf nTrtltal (Ktlnn of a iinini t>n<l. lucli u of UorMKbestnnt 
N. TIw ■<!■ nr Uin Hinii derctiiplnt. II'B clcfflgathm b(«lnnln^' willi tlic ■.■khI liiric- 
nndg.non ftillovorl hj tlie othen In inoccinlati. NL A itur'i gmwlli of HitnorlmiBnt, 
cnwnttl wllh > Mnnlnul hod: a, (ctnlvn bj (ba biiil-«cida of tlie |>rgriuua jt»r: 
b, *»n left H Uie Ikllvn lmr->(al1a; c, kxllUrr bD<l>. 

riQ. HC BninrhilniirniilcrmllBiUUrDnrthnlJUe 

FIU. er. Lear-bud uiukr Uie pcllolo of Ihc F1iu»-tra«> 


73. fleshy Buds. BuHn are peculiar buds of certain herba- 
ceous plants, with fleshy scales, and often of a more permanent 
character. Their nature and economy maj- most conveniently 
be illustrated under subsequent sections. Usually bulbs are 
subterranean or partly so. But small bulbs {Buiblets, 123) regu- 
larly appear in the axil of nearly all the leaves of certain common 
Lilies, being obviously ordinary axillary- buds, under certain 
modifications. They become detached at maturity, fall to the 
git>und, produce roots, and grow as independent plants ; and 
their flesh}- scales are storehouses of nourishment for the early 
support of this independent growth. 

74. Bud-propagation is a normal mode of reproduction in cases 
like the above, the spontaneously detached buiblets or buds 
establishing themselves as progeny. In several species of 
Allium (Onions and Leeks), such buiblets usurp the place of 
flower-buds, making the analogy seem closer. Stems or branches 
which habitually root in the soil, or along its surface, equally 
propagate or divide into new individuals, becoming distinct by 
the perishing of the older connecting parts, or by breaking away 
from them. Propagation by cuttings is an acceleration or exten- 
sion of this same natural operation. The cutting is a portion of 
stem bearing one or more buds, which, through the faculty of 
the stem to strike root, is made to gi*ow inde|)endently. In 
grafting^ such a cutting, and in budding a bud only, with a small 
portion of wood and bark, is transferred to the stem of another 
plant of the same or of some related species, and made to grow 
there, uniting its wood and bark with those of the stock, and so 
becoming a limb or branch, in place of striking ix>ot into the soil 
and becoming a separate plant. The horticultural advantage of 
bud-propagation is, that the offsets or new individuals share 
in aU the peculiarities of the parent as completely as if still 
branches of that tree. In propagation by seeil, the special 
peculiarities or excellencies of individuals or varieties may not, 
and in some measure probably will not, be reproduced. 

75. Normal or Begnlar Bads^ as to position, are either terminal 
or axiUary^ as already stated. (15.) They are single, that is, 
one bud normally occupies the apex of a stem or branch, and 
appears, or usually may appear, in the axil of (or upjKjr angle 
formed with the stem by) any well-developed leaf. In these 
positions, buds are so usual, or so capable of appearing, tliat 
they are commonly regarded as potential when not actually 
present. The potentiality may be manifested by the actual 
development of these buds in shrubs or trees after the lapse of 
years. (84.) The terminal leaf-bud is to continue the axis it 


stirmouDts : axillarj' &wl any other lateral leaf-buds are to be- 
come branebes. But even of buds which actually appear a large 
proportiuD do not grow. When a termiaal bud is formL-d (as 
Fig- 81, 85, 91), this is commonly tbe strongest, or among the 
stronger. But iu many cases it babitunlly or commonly fails to 
a|}i>ear. In the Elm (with leaves and therefore buds alternate), 
the bud axillary to llie last leaf of Ihc season takes its plaee. 
In the common Lilac, a pair of buds, which wero in the axib of 
the uppermost of the (opimsilc) leaves, seem to replace the 
terminal bud, which seldom <leveloi>s. (Fig. 8C.) When all tbe 
wgiilar buds make their appearance, and llic leaves are opposite, 
Uie stem will l>c crowned with tlic terminal bud, having an axil- 
lary bud ou each side of it, (Fig, 8S.) 

7G. Arcessur; Bpda. These are, as it were, multiplications of 
the regular axillary bud, giimg rise to two, three, or more, instead 

* of one ; in some cases situated one above lui- 
other {wperposed), in others jilaceil side by 
side {nJlateral). In tlie latter case, which 
occurs occasionally in the Hawthorn, in cer- 
t4un Willows, in tlie Maples (Fig. 88), 4c., 
the axillarj' bud seems to divide into three, 
or itself to give rise to a lateral bud on each 
si<lc. On some shoots of the Tartarean 
Honeysuckle (Fig. 90) from three to six bnds 
apitear in each axil, one altove another, the 
lower being snccessively the stronger and 
eailier producetl ; and the one immeiliatety 
in the axil, therefore, grows in pi-eference : 
oecHsionally two or more of them gi-ow. and 
8U|]ei'poscd accessoiy branches n-sult. It is 
much tlie same in Aristolochia Siphu, except 
that the upiwrmost bud is there sti'ongest. 

FIQ. se. Brueb of Rsl M*t^e. at 
Mc liy (Ida. 

riQ. 89. PhwenriilintnEhiir UioB 
kii»Tber; a, the teaf ksi ; h, prnpnt iiill1iii7 liuil; «, 

Fin. on. Pari <>f ahrUNli iiT Turutcui Uuncyi 
tu|ierp(iMi) In Ihe oiJI ormclj RuiT. 

mlilille btwlng tri|ila ullUry biiili. plknd 
liiiil* plmva nne sbors 


So it is in the Butternut (Fig. 89) , where the trae axillary bud 
is minute and usually remains latent, while the accessory ones 
are considerably remote, and the uppermost, which is much the 
strongest, is far out of the axil : this usually develops, and gives 
rise to an extra-axillary branch. 

77. Adrentitioos Buds are such as are abnormal and irregular, 
being produced without order and from any part of the stem, or 
even from roots. The latter, like the internodes of a stem, 
although normally destitute of buds, do produce them notwith- 
standing in certain cases, esi)ecially when wounded, and in some 
plants (feuch as Blackberries) so freely that gardeners propagate 
them b}- root-cuttings. The stems share this tendency; and 
buds are apt to break out on the sides of trunks, especially when 
wounded or pollarded, or to spring from new tissues produced 
on cut surfaces, especially where the bark and wood join. Even 
leaves may develop adventitious buds, and then be used for 
propagation. In Bryophyllum, such buds, followed by rootlets, 
are freely pixxluced on the mai-gins of the blade or of its leaflets. 
In Begonia, a leaf, used as a cutting, will root from the base of 
the petiole stuck in the soil, and produce buds on the blade, at 
the junction with the petiole, or elsewhere. 

Section III. Of the Stem. 

§ 1. General Characteristics and Growth. 

78. The Stem is the ascending axis, or that iK)rtion of the 
trunk which in the embr}0 grows in an opposite direction from 
the root, seeking the light, and exposing itself as much as pos- 
sible to the air. All phienogamous plants ix)ssess stems. ^ In 
those which, in botanical descriptions, are said to be acaulescenf^ 
or stemless^ it is either very short, or concealed beneath the 
ground. Although the stem always takes an ascending direction 
at the commencement of its growth, it does not unifonnly retain 
it ; but sometimes trails along the surface of the giound, or 
bunx)ws beneath it, sending up branches, flower-stalks, or leaves 
into the air. The common idea, that all the subteiranean jwrtion 
of a plant belongs to the root, is incorrect. Equally incoiTcct is 
the common expression that plants spring from the root. Roots 
spring from the stem, not the stem from the root. (21, 24, 37, 44.) 

1 There are, however, reduced forms in which there is no distinction of 
axis and foliage ; but most of these are clearly leafless rather»than stemless, 
and not even in Lemna and Wolffla can the stem be said to be wanting. 



79. While the root normally gives birth to no other orgai», 
t)ut itself |Jorforma those Ajnetioiia which pertain to the relation§ 
of the vegetohle with the soil, — binding it to the earth and 
alisorhing Qonrisbing materials from it. — the aerial fiiuetioue of 
vegetation are chieflj' earned on, not so much by the 8l«m it- 
self as by h distinct set of organs whieb it bears, namely, the 
leaves. Hence, the production of leaves is one of the chamc- 
teristics of the stem. These are produced only at certain dednito 
and symmetrically arranged rioints, called nodes. (13, 23.) 

80. Derclopmeut aad Stmctare. In a bud or undeveloped st«n. 
the nodes are in contact or close proximitj. In the develoj)- 
incnt, growth in tongtii takes place in such manner as to carry 
these apart more or less, according to the degree of elongation, 
that is, tJie inlernodes (13) elongate. The oi'dcr of development 
is fh>m below upward, the lowest intemode first lengthening, 
the others in regular succession. Each completes its growth, 
with more or less rapidity, although the length attained varies 
greatly in different stems, in different parts of the same stem, 
and under different conditions. llnUlte the root, in nliich the 
elongation of formed ports is verj- soon finished and tlicrcfore 
only the tip is perceptibly growing, intcmodes go on growing 
throughout, and several formed intcrnodcs may be grawiiig 
simultaneously, thus producing elongation tliroughout a consid- 
erable extent of stem and with considerable rapidity. But each 
intemode grows independently. Some parts of an intemode 
may lengthen faster or continue in growth longer than others ; 
this is usually the upper portion, at least in long intemodes and 
when ever}- i«irt is equally e.tjwsed to light. 

81. The development of a sl«m from a bud is wholly Uke that 
from the embryo, and has already been doscrilied in Chap. li- 
lt exhibits similar variations as to rapiditj' and \'igor: dependent 
npon the constitution of the bud, — which, like tlie plumule in 
the seed or seedling, may be cither latent or much develo|)ed 
before growth begins, — also upon the amoimt of nourishment 
prorided. Strong buds commonly have tlieir parts, or some of 
them, ready forminl in miniature, and a store of elalwrated nour- 
ishment in the parent stem to draw upon. Those well-developed 
buds which in many of our shrubs and trees crown the apex or 
occupy the axils of stem and branches early in tlie preceding 
summer (as in ilagnolia. Fig. 81. Horseehestnut. Fig. S3, and 
Hickory, Fig. 91) often exhibit the whole plan and amount of 
the next yeai's growth ; the nodes, the leaves they bear, and 
sometimes the blossoms being already formed, and only requiring 
the elongation of the intemodes for their full expansion. As 





the bud is well supplied with nourishment in spring by the stem 
on which it rests, its axis elongates rapidl}' ; and although the 
growth commences with the lowest internode, 3'et 
the second, third, and fourth intemodes Ina}- 
begin to lengthen long before the first has attained 
its full growth. Such very strong buds are usually 
terminal ; but sometimes, as in Lilac (Fig. 86)^ the}' V^J 
are the uppermost axlllar}', which take the place of Vr»Sr/ 
a suppressed or abortive terminal bud. 

82. Such wood}' stems, develoi>ed from a strong 

bud, and terminated at the close of the season's 


growth b}' a similar bud, may be continued from 

3'ear to year in an unbroken scries. A set of narrow 

rings on the bark (Fig. 85 a) commonly marks 

the limit of each year's growth. These are the 

scars left by the fall of the scales of the bud ; and 

these, in the Horsechestnut, and in other trees with 

large scaly buds, may be traced back on the stem 

for a series of years, growing fainter with age, 

until the}' are at length obliterated by the action of 

the weather and the distention caused by the increase *^ 

of the stem in diameter. The same is the case with the more 

conspicuous Leaf-tcan^ or marks on the bark left by the separation 

of the leaf-stalk, which are for a long time conspicuous ou the 

shoots of the Horsechestnut (Fig. 85 li) , the Magnolia (Fig. 81), 

and Hickor}', Fig. 91. 

83. Ramillcatioii. Branches (14-16) are secondary stems 
developed from a primary one, or tertiar}' ones from these, and 
so on. Ultimate or small ramifications of latest order are some- 
times called Branchlets. The terminal bud continues the stem 
or axis which bears it. Lateral buds give rise to branches.* 
As the normal lateral buds are axillar}' (75), so are normal 
branches. The s}'mmetr}' or arrangement of branches, being 
that of the buds from which they are developed, is fixed by and 
follows that of the leaves. When the leaves are alternate, the 

1 Dichotomy or forking, the division of an apex into two, although of com- 
mon occurrence in the lower cryptogamous plants, occurs so rarely and 
exceptionally, if at all, in phaenogamous plants that it may here be left 
out of Tiew. 

In phcnogamous plants only the ramification of axes should take the 
name of branches. That is, roots and stems branch ; and the term may 
without confusion be extended to hairs and all Tricuomes (383) when com- 
pound, but not to leares and their modifications. 

FIG. 91. End of a Hickory branch (Garya alba), with a strong terminal and smallw 
axOtaiy badu 



ItrsQchea will be alternate ; when the leaves are oppodte, 1 
the bads develop regTilariv, the branches will be 0|>posite, &c. 
This hol<Is in fact siiltlcieittly to determine and eseni{ilirt' the 
])lan of ramification ; but. if entirely carried out, there would be 
as many brauches as knves. This conld rarely if ever Iw, even 
ill primary ramification. 

84. Xon-dfTelopmeat ftf Bads. Some of the buds arc latent or 
merely [xttential, that is, do not make their apiiearance : of those 
which do apiieur only a part actually grow into brandies ; and 
of theae some are apt to perish at an early stage. In our tives, 
most of the lateral buds geueraily remain dormant for the first 
season : they appear in the asils of the leaves early in Burainer, 
but do not grow into branches nntil the following spring ; and 
even then only a [wirt of them grow. Sometimes the failure 
occurs without appreciable order : but it often is nearly uniform 
in each species. Thus, when tlie leaves are opposite, there are 
usually three buds at the apex of a branch ; namely, the terminal, 
and one in the axil of ench leaf; but it seldom hap|>ens tliat all 
throe develop at the same lime. Sometimes the terminal bud 
continues the branch, the two lateral generally remaining latent, 
as in the llorsecheatnut (Fig. 85) ; sometimes the temiiual one 
fails, and the lateral ones grow, when the stem annually becomes 
two-forkeil, as in the Lilac, Fig. 6€>. The undevelo|>ed buds 
do not necessarily perish, but are ready to be called into action 
in case the others arc checkeil. When the stronger buds are 
destroycfl, some that would else remain dormant develop in their 
Bte^d, incited by the abundance of nourishment, which the for- 
mer would have monopolized. In this manner our trees are soon 
reciothed «itli verdure, after their tender foliage and branches 
have lieeii killed by a late vemol ftoat. or consumed by insects. 
And buds which have remained latent for several years occasion- 
ally sliool fortli into branches from the sides of old stems, 
especially in certain trees. 

85. Most )>ranchcs springing from old trunks, however, as in 
Willows and Pu|iiars. esi^cially when wounde<i or i>ollarde<i. 
»riginnti« fVom adventitious buds (77), which occur without 
order. So also when accessor}' buds (76) develop into branches, 
normal symmetry is more or less di8turl)ed, as by contiguous 
slioola standing direct!}" over each other in Tartarean lioney- 
sucklc, or by a branch far out of the axil in Walnuts (Fig. 89) 
and Honey-Locust, Fig. 'id. 

8G. Excnrrent and Delli|uegcent Stems. Sometimes the primary- 
axis is prolongeil without interruption, even through the whole 
life of a tree (unless nceidcntaily destroyed), liy tiie continued 


evolution of a terminal bud, or by some upper strong bud which 
equall}' becomes a leader, — forming an undivided main trunk, 
from which lateral branches proceed ; as in most Fir-trees. 
Such a trunk is said to be excurrent. In other cases, the main 
stem is arrested, sooner or later, either by flowering, by the 
failure of the terminal shoot, or by the more vigorous develop- 
ment of some of the lateral buds ; and thus the trunk is dissolved 
into branches, or is deliquescent^ as in the White Elm and most of 
our deciduous-leaved trees. The first naturally gives rise to coni- 
cal or spire-sha[)ed trees ; the second, to rounded or spreading 
forms. As stems extend upward and evolve new branches, those 
near the base, being overshadowed, are apt to perish, and thus 
the trunk becomes naked below. This strikingl}* occurs in the 
excurrent trunks of Firs and Pines, grown in forest, which seem 
to have been branchless to a great height. But the knots in 
the centre of the wood are the bases of branches, which have 
long since perished, and have been covered with a great number 
of annual layers of wood, forming the clear stuff o^ the trunk. 

87. Definite and Indefinite Annual Growth of Branches* In 
man}' of our trees and shrubs, esjKJciall}* those with scaly buds, 
the whole ^•ear's growth (except on certain vigorous shoots) is 
either already laid down rudimentally in the bud, or else is early 
formed, and the development is complete<l long before the end 
of summer ; when the shoot is crowned with a vigorous terminal 
bud, as in the Ilorsechestnut (Fig. 85) and Magnolia (Fig. 81), 
or with the uppermost axillary buds, as in the Lilac (Fig. 86) 
and Elm. Such definite shoots do not die down at all the follow- 
ing winter, but grow on directl}', the next spring, from these 
terminal or upper buds, which are generally more vigorous than 
those lower down. In other cases, on the contrary, the brandies 
grow onward indefinitely^ until arrested by the cold of autumn : 
the buds at or near their summit are consequently young and 
unmatured, or at least the lower and older axillan' buds are 
more vigorous, and alone develop into branches the next spring ; 
the later-formed upper portion most commonly perishing from 
the apex downwartl for a certain length in the winter. The 
Rose and Raspbeny, and among trees the Sumac and Iloney- 
Locust, are good illustrations of this sort; and so arc most 
perennial herbs, their stems dying down to or beneatli the sur- 
face of the ground, where the persistent base is charged with 
vigorous buds, well protected by the ground, for the next year's 

88. Many of the details and applications of ramification, of 
most importance in morphology- and descriptive botan}', relate 




to anthotaxy or inflorescence (Chap. V.), which has its ow^ 
termiuology. But some ol" its ti^rms may be conveniently 
employed iu the description of ramiScatioii uiiuotiiieeti^ with 


§ 2. Forms or Stems axd Bran-ciies. 

89, On tlie size and duration of the stem the oldest and most 
ohviouB division of plauta is founded, namely, into Herbs, 
Shrubs, and Trees. 

DO. Herbs are [ilants In which the stem does not become 
wood,v and ijorsist^nt, but dies annually or after flowering, down 
to the ground at least. The dittcrence Itetween annual, bimnial, 
and pgrcmiiai herbs has already been point^Ml out in the cha|>ter 
on the root (aO-57), and the gradations between them indicat«d. 
Herbs pass into shrubs and shrubs into trees through every gra- 
dation. The folbwing {liidnitions ai-e therefore only general ; — 

91. I'ntffrslirutw, or Suffrulicoie plants, are woody plants of 
humble stature, their stems rising little above the surlhee. If 
less decidedly wikx1_\', they are termed. SuffnaetcetU. 

1)2. Shmbs are woody plants, with stvms branched fivm or 
near tbci ground, and less than five limes the helglit of a mau. 
A shnib which approaches a tree in size, or imitates it in aspect, 
is Rni<l to be Arboretcimt. 

93. Tre«e are wootly plants with single trunks, which attain 
at least four or five times the human stature. Yet the name of 
tree is not to be denied to a woody plant having a single and 
stout trunk of less altitude : and those which grow in a bushy 
manner, sending up a cluster of stems frwm the ground to the 
height of tliirtj- feet or more, may still lie called shrubB. 

04. The erect |)osition, elevation above the soil, and self-sup- 
port, are normal conditions ofthe stem, but are far (Vom universal. 
And certain kinds of stem or branches arc sufficiently poouliar 
to have wceived «nl>staiiH\'e names ; other equally peculiar forms 
have no sjK'Cial names. There are, moreover, certain orgaps 
(sudi as spines and tendrils) which are commonly homologous 
(12) wiUi stems, but not always. Two kinds of erect stems 
lia\o special names in descriptive botany. 

95. Culm is a name applied to tlie peculiar closed-jointed stem 
ofGrasses and Sedges, wliether herliaeeous. as in most Grasses, 
or woody or arborescent, as in the Bamlioo. 

9G. Candeic is the name technically applied to the trunk of 
Palms (Kig. I2C), Troe-Fcma, and tlie like, consisting of a 
commonly simple column, the surface beset with scales, — 


bases of former leaf-stalks, — or marked by sears, left by their 
fall. This name was U8e<l by botanists anterior to Linnaeus for 
an}' tree-trunk, but is now used for the [>eculiar stems above- 
mentioned ; also for the persistent base of a stem, otherwise 
annual, whieh throws up fi-esh herbaceous stems or stalks from 
3'ear to 3'ear. Such short and enduring stems, hemg usuall}* 
near the ground or under it, were commonl}* mistaken for roots. 
The old English name of Stock is sometimes used in botanical 
description for all short and enduring stems of this sort, 
whether rising somewhat above or concealed beneath the surface 
of the soil. 

97. A Scape is a stem or branch which rises from beneath or 
near the surface of the ground and bears flowers, but no proper 
foliage. It therefore belongs to inflorescence. (2().").) Scapes 
usually spring from some one of the subterranean fonns of stem. 

98. Of stems which do not stand upright in the air there are 
various modifications and gradations. 

99. Scandent or Climbing Steins are those which rise b}' 
attaching themselves to some extraneous support. This is 
effected in various wa^'s ; in some by the action of the stem 
itself, in others by that of organs which it bears. ^ 

100. Tolnble or Twining Stems, or Twiners^ are those which 
ascend b}- coiling round a support, which must accordingly* be 
comparatively' slender, or at least not too large. Some ascend 
b}' coiUng ''with the sun" (that is, from right to left of the 
obsen'er viewing the coil from the outside ^) , as the Hop ; more. 

1 See Darwin, The Movements and Habits of Climbing Plants, I^ndon 
and New York, 1875. Also the earlier paper on the subject in Journal of 
the Linnean Society, ix. 1865. 

Note that in North America climbing plants in general arc in popular 
language called Vines («. g. Hop- Vine, Grape- Vine, Squash- Vine, &c.), a 
name which properly belongs to Vitis only. 

« DertroraeAud Sinistrone, i. e. to the right or to the left, arc almost indis- 
pensable terms, but there is an ambiguity and discrepancy in their use. 
Darwin (in Climbing Plants, above referred to) seeks to avoid this by usually 
employing the terms " with the sun," and "against the sun," phrases which 
would be unmanageable in terminology. The writer (in Amer. Jour. Sci. 
ser. 3, xiii. 391) suggested Eutropic for the former, Antltropic for the latter, 
to be used in case it is preferred to evade rather than to encounter the 
ambiguity. Probably the terms dextrorse and sinistrorse, or right and left, 
will continue in use, as most natural and convenient. Now, in the first 
place, it should be understood that a plant, or at least a plant's axis, having 
no front and back, can have no right and left of its own. These relations 
of direction must refer to the right and left of an observer. All depends, 
accordingly, upon the position which the viewing observer is supposed to 
occupy when he predicates the direction of the turns of a helix or of the over- 
lapping of the parts of a bud. Linnaeus supposed the observer to view the 



by coiling' in the ojijioaite direction, as the Bean (Pbaseolua), 
the woo(i_v AristuliK-hLi Sipho, the Morning Glorj- (Fig. 91') atjil 
. otlier CoiiYolvu- 
, lucwL". Tlic Uoil- 
', a leafless par- 
asitic plant of the 
r fnniily. not only gains suppoit by coiling 
on the steins of other plants, but by attachment, 
through the devcloprnent of siiokcr-lilie discs, 
along the whole conligiiouu surface. (Fig. 77.) 
The various attions througli which plants climb, 
and the attendant phenomena, are physiological, 
ami nnl! bo treated in the second part of this 
Text Utxik. The most complete and satisfac- 
tory discussion of the subject, of a readable sort, 
ts that of Darwin's volume, referred to in a 
preceding not*. 

101. L«if-CIimbers are those in which support is gained by the ' 
action, not of the stem itself, but of the leaves it bears ; in most 
by the coiling or clasping of petioles, as in Clematis, Maurandia. 
Tropicolum, and Koluniim jasrainoides (Fig. 235) ; in some by 
the inciir\-ation of leaf-blades or portions of them, as in Adlu- 
mia ; or by an extension of the midrib into a hook or short Uin- 
(Iril, as in Gloriosa; or by the transformation of some of the 
blades of a coni|X)und leaf into boolcs or tendrils, as in Cobsa 
anil the Pen. 

Vr2. TendrlLCIImherB (Fig. !)2-9.i) arc those in which the 
prehension is by a tendril, u slender filifonn ho<iy, either simple 
or branched, specially adapted to the puriiose. and capable of 
coiling, either to sccnrc a hold, or to draw the stem up to the 

oiiit or cirfle (rom the iiieiilc ; Mohl, Pnt'm, Brnun, and the I>eCftn(lD]1t« 
adupt IliiB, and llir Inttcr hnisE on il. Such Ruilmrity slmulil In- decisive. 
if nimmnn aiia^ ami popular tense went alnn^ wiili ii. Bui aome ol tlie 
bolanists following Linnteiu adopted the ivrerse view : and to the present to Ueiilliani Hnd Hooker. IMrw in. liiclikr, ntid in pnrtG. Hcnilow, 
it wa» wi natural to Tiew ||,^ ^,{] tmm the ouiiiJe that ire witliout e 
adopted this position anil mode- of cupresainn. A righi-liand coil, or one 
Inrning to tlic ri^lit, with ua. is one the (ilrna of vhii:h pMs from tlie left to 
riglit of a bf Btnmler wlio confronts llie coil. 

nion lerew i* called a right-liancieil screw. and tliat tliL- right bnok at a river 
i> tliat to tlie right of the person who follows the eoane of ilic si 
natural is this, that even on a map or plate, which has foee and haek, and 
therefore a riglit iind left ol lis own. the Hgiire* oecupyin^ tti> right nr left 
portions are andcnUHHl to be those whith are toward thu right or llit- left 
hand of the ohserrer wlii> stati<la l)etore it. 
. Dulnn^cly (wtnl 


support. In certain tendrils the attachment to the 8up|»rt is by 
a 8ucker-like disc at the ai>cx, as in the Vii'giiiia C'reej^er or 
Amivclopsis, Fig. 94. 

103. Koot-€Ilnben are those in vhich the stems [iroducc aerial 
rootlets ((i'J*). which fix themselves to a supiMiting surface 
along which the stem creeps or ascends. In this wny Trumiwt 
Cree|)er (Tecoma radtcans) , Ivy, and Poison Ivy (Uhns Toxi- 
codendron) climb extensively. 

1U4. Stems or branches which neither climb nor stand upright 
may have their direction or habit of growth expressed by certain 
adjective terms ; such as 

AKenditiff or Asturgent, when they rise obliquely upward ; 

Rtclining, when from an ascending or erect base the upper 
part recur\'es and trails ; 

Decumbent, when trailing along the ground, but with apex 
assurgent ; 

Proevmbent or Prvttraie, when lying at length n|K»n the ground ; 

ReptTtt or Creeping, when growing prostrate on the ground 
and rooting as they grow. Abo a[>plic<l to similar stems grow- 
ing under, as well as upon the surface of the soil, as in Couch- 
Grass and Mint, Fig. 99. 

10a. A Sncker {tiureulus) is an ascending stem rising fVom a 
snbten-anean creeping base. The Kose and Ra8|»berry multiply 
freely by suckers. Such plants are easiest to propagate *■ by 

IOC. A Stolon is a prostrate or reclined branch which strikes 
root at the tip, and then deielops an ascending growth, which 
becomes an independent plant. 

107. An (HRtet is a short stolon or a short sncker. Ilouseleek 
(Fig. 91') offers a familiar 
example. By offsets, some 
herbs, otherwise annuals, are 
continued from year to jear in 
a vegetative progeny (Lobelia 
canlinalis, &c.), and peren-f 
nints may thus establish colo- 
nies arounda imrent individual. 

108. A Btuner (Flagellum) 
is a Hliform or \ery slender »i' 
stolon, naked and tendril-like exce[>t at tip, where it roots, 
develops a bud, and so a new plant. The Strawberrj' tlimishes 
the most familiar example. 

FIG. «l*. BooMlMk (ScmpenlTaai (Ktnruiiil vHh ufflwt*. 



109. The two following are organs whith maj- be of axial 
nuture, or may not. This may ordinarily be determined by posi- 
tion. Any direct continuation of 
stem or branch muet be of axial 
nature, that la, of the nature of stem ; 
and the same is true of whatever 
primarily develops in the axil of a 
leaf. Conversely, whatever subtends 
a kteral axis or branch may be taken 
for a leaf or foliar production, being 
in the place of such. 

110. A. Tendril, a thread -slia|>ed 
f and leafless Ixxly, capable of coiling 
L spirally, and used for climbing (102), 
is homolt^ous with stem in Grape- 
3s (Fig, 1)2) ; for the uppermost 
t«i)dril is seen to be a direct continu- 
ation of the stem. The small bud 
which a[i])Oars in the axil of the 
"V^^-vV jl f'^J\ upjwnnost leaf will in its growth 
V -'' \^'=^^^^ proiluee another intcrnode and leaf, 
^~^'7iy''^\J[.Ij?\j or some species more than one, 
n but will terminate in a simikir 

tendril : the present terminal tendril will have thou become 
lateral and opiwsite the leaf, like the three in the lower part of 

FIO. !I2. Eiiil lit a I 
FIG. W. A pnriliHioi 
Plti. »i. Kiitli or Ibv 

t (lia aisiw-rliie, Willi Jiuuiig leiHlrltK: k aylD|»<lluI 
1 of Ampelopuls <]iiln<|uefoll>, «r rirglnlk Crc«iwr, vUh 
eiilBTgvil. (binrlnj; the ei|«iiJoil tlim ur ^lllc• bjr olikli 


tlie figure.' The teDdrils of Virginia Creeper (Fig, 93) are of the 
same nature and ixtsition. But, instead of laying hold hy a coiling 
of the tip, when it has reached any solid surface, such as a wall 
or tree-trunk, the tip expands into an adlieaive disc, which forms 
a secure attachment. (Kig. 9-1.) In a related plant, Vitis{Ci8sus) 
tricuspidata of Japan, these disks terminate the branches of ver}' 
short tendiils: consoquentlj- the shoots as they grow are at 
once applied closely and seeuretl finnly to tl)c surface of the sup- 
port, — an admirable adaptatioD for climbing walls and trunks. 

Ill, Tlic simple tendril of a I'sssion- 
flower. being in the nxi! of a loaf (lliat 
is in the position of a branch), is also of 
axial nature : it is a leafless and sinij)lc 
branch, com[x>sed of one long and slen- 
der intemocle, devoted to the purjiose of 
climbing. Fig. Ho shows in all stages the 
admirably active tendrils of Passiflora 
eioyoides. This is a KIcxican si>ecies, 
remarkable for the rapiilily and freedom 
with which the tendiiU move. The lowest 
tondiil in the flgure is attached and 
coiled : the next is IVee and coiled in 
one helix : the tliird is outstretched 
and seeking a support. For tendrils 
which arc not homologous with stems, 
see Sect. IV. 22S. 
112. A^IseorThom (Fig. 96, 97) is usually a branch or 
the termination of a stem or branch, indurated, leafless, and 
attenuated to a point. The nature of spines is manifest in the 
Ilawthoro (Fig. 97), not only by their position in the axil of a 
leaf, but often by producing imperfect leaves nnd buds. And 
in the Sloe, Pear, &c., many of the stinted branches lieconie 
tpinoit or spinricent at the apex, tapering off gradually into a 
rigid and leafless iwint, thus exhibiting cveiy gradation lietween a 
spine and an ordinaiy branch, Tliese spinoso l)ianches are less 

1 Tbil fomu what ia called a S^m/mliHw or Sniipodinl Urm. wlik'li ia mnr- 
phulogicallj made up of a acrira of supprponod branches, (See Chapter V. 
281,282.) Ia contradistinction, a stem formed hy tlie continued (leTeliipnient 
of a terminal bud is ifimo/iodiai or a MonoiKdivm. Fig. 05 ia an example. 

VIO. M. \/»fi sbcrat nf Purirlnra alvTotilea, of H«ilco, wltli flied ami coUed, (ne 
and foU grown, awl tbrmlug tsnililli. 



liable to appear on the cultivated tree, when dul}' cared for, such 
branches being thrown mostly into more vigorous growth. In 

the Hawthorn, the spines 
spring from the main axillary- 
bud, while accessory buds 
(7G), one on each side, ap- 
pear, and grow the next sea- 
son into oi*dinary branches. 
In the Honey-Locust, it is 
the uppermost of several ac- 
cessory buds, placed far above 
the axil, that develops into 
the thorn (Fig. 96) . Here the 
spine itself usually branches, 
and sometimes becomes ex- 
tremely compound. 

113. For spines which are 
homologous with leaves, or 
parts of a leaf, see Sect. IV. 
227". Prickles^ such as those 
of Brambles and Roses, are 
sui^erficial outgi*owths from 
thci bark, of a different nature 
(383), and of small mor^^ho- 
logical signification. 

114. Subterranean Stems ai*e hardly less diverse than the 
aerial. The}* are classed as Rhizomes, Tubers, Corms, and 
Bulbs, the forms passing one into another b}* gradations. 

115. Rhizoma {Rhizome^ or in English Rootstock) is a gen- 
eral name for any horizonal or oblique perennial stem, which lies 
on the ground or is buried beneath its surface. It sends off 
roots of a fibrous or slender sort wherever it rests on or is cov- 
ei-ed by the soil, and usually produces from its apex some kind 
of aeinal stem, either leafy or as a flower-stalk (scape^ 97), 
which rises into the air and light. Before morphology was 
understood, rootstocks were called creeping roots^ scaly roots, &c. 
Some are slender, such as those of Mints (Fig. 09), of most 
Sedges (Fig. 98), and of Couch-Grass . Their cauline nature is 
evident from their stnicture and apiwarance ; their nodes and 
intemodes are well marked, the former l)earing leaves reduced to 

FIG. 96. Branching thorn of tlie Honey- IxvuRt '01«Utf»rliia). an indurated branch 
develop<»<l fh>m an acce«w»ry bud j»ro«luce<l al»ovo the axil. a. Three hmU under the 
baiie of the leaf-Htalk. brought to view In aMvtlon of the ntem and leaf-ntalk below. 

FIQ. 97. Tliom of the Cockspur Tliom. developed from the central of three axillary 
budB ; one of the lateral buds is »een at itM base. 



scales ; and the advancing ap>ex rises at length into an ordinar}' 
stem, while the opposite and older end gradually dies away. A 

bud forms in the axil of each scale-like leaf, or 
in some of them ; roots proceed from the nodes 
in preference ; the destruction of the ascending 
stem only brings these buds into activity ; and 
the cutting or tearing of the rootstock into 
pieces by the hoc or plough merely hastens the 
establishment of as man}' new plants, each with 
roots, bud, and a small store of nourishment 
read}' provided. It is this which makes Couch- 
Grass or Quick-Grass (Triticum repens) very 

troublesome to the agriculturist ; and the Nut-Grass (Cyperus 
rotundus, var. Hydra) of the Southern Atlantic States is even 

more so, portions of its rootstock being tuberiferous, t. e, en- 
larged into a tuber which contains a supply of 
concentrated nourishment to feed the growth. 

116. Thickened rootstocks are common; 
nourishing matter, elaborated in the leaves 
above, l>eing accumulated in them, just as 
it is in thickened roots, and for the same pur- 
pose. (53-55.) Such are the so-called roots of Sweet-Flag, of 
Ginger, of Iris or Flower-de-Luce (Fig. 216), of Bloodroot, of 
Solomon's Seal (Fig. 100), &c. These grow after the manner 
of ordinary' stems, advancing from year to year by the annual 
development of a bud at the apex, and emitting roots fi-om the 
under side or the whole surface. Thus established, the older 

FIG. 98. Slender rbizoma of Carex arenaria, of Earope, which binds sliiftiiifi^ sands 
of the sea-tibore. 

FIG. 99. Rootstocks, or creeping sabterranean branches, of the Peppermint. 

FIG. 99>. A pie<*e of the rootstock of the Peppermint, enlarged, with its node or Joint, 
and two axillary badB ready to grow. 


[iDi-tions die and detay as corrc9|K)uding additiuna are i 

tlie opposite growing extremity. Eacli year's growth is often 

marked conapicuoiisly, sunf^tinios by a stmng contraction where 
the inteituptioutook plate, as in eertainapcciesof Iii8(l'1g. 216); 
or hy the circular im- 
pressed scar (likened to 
the iiiipressiou ora eeal) 
' in Solomon's SeiU ; tliis 
being the place where 
iLe annual aerial stem, 
bearing the vegetation, 
sei>arated in autumn 
Trom tlie |K'reniiial rhi- 
zoma. The numerous 
slender lines encirclingthe i\X)tstoek are the SL'ai-a left aft^r the 
decay of tha scale-like leaves or hiid-scalea, such as are eeea at 
the young and growing end of the rootstuck. 
The rootstock of Dipliylk-iu, of lUe Alleghany 
Mountains (Fig. 101), is similar; but the 
yearly growths are so exceedingly short that 
they !>eeome vortical, the bud of each year 
is close to the stalk of the year preceding, 
Lud the sears marking previous gruwtlia are 
ill contact.* Trillium makes a short and 
iMustly vertical rootstock. which, when it 
rcnmina simple and ilies away promptly 
liclow (as ill Fig. 102), comes nearly wiltUn 
tlie ilefinition of a corm. But in several 

1 TLe ruutitiK'k in Pulytsuiiatuni and Diphyllciu is > lympinliiim (UO.xorrl, 
Hu! ICTiniaal bud dcvelupiiig jiearly the (^vrih above ptiund niitl perishing 

« Kan from n-Lich tbe l«t<r hu Kpmlol Id 

FIO. IDl. Khluimiinf DI|>1iy1lrliicrinniNt.*h 
the HTnnth: n, tha htui: b, bua oT 111 

Fill, liri Sl»wl B1..1 roalig tuoUtuck 



species, and in older individaala, it is longer, often oblique, and 

branching, and bears the Bears from which the annual aerial 

growths bare separated.^ Nyinphfea odorata, 

the sweet-scented white Water Lily, grows by 

verj' long, stout, and simple rootstocks. In 

N. tuberosa the sides of the rootstock produce 

short lateral branches or tubers. 

117, A Tuber may be morphologically char- 
acterized as a short thickened rhizoma on a 
Blender base, or a rootstock some portion of 
which — mostlya terminal portion and involv- 
ing several nodes— is thickened by the dejx)- 
sition of DourishiDg matter. A potato and a 
Jerusalem artichoke are typical examples 
(Fig. 104-lOT) ; and the difference between 
these subterranean branches and the roots which they may bear 
is verj' obnous. Their eyes are axillary buds ; the leaves which 
subtend them are plainly dis- •*• "» 

cemible, in the form of short and 
closely appresseil scales. In the 
attempt, occasionally seen, to 
form asillarj' tubers above- 
ground by the Potato-plant, the 
leal^' nature of the scales is 
evidenced. (Fig. 
10o.)By heaping 
the soil arc 
the stems, the 
iferoiis branches 
may be 

creased. The number of nodes and internodes involved in a 
tuber may be many or few. There is one instance of what may 

in RUlumni to be renewed bj an axillary bu<l, which makre its Bubteiraneaa 
growth and (he mdimpnt* af tlie ncrial in tnrly sunimor. 

' Thi« rhiioma is a monnpodium, beiiiR coniinued yoar by year by the 
lerminsl bud, anil the aerial slum ur etemi aent up in spring, bearing the 
whorl of leave* and bloMom, are axillary branchce. 

Fia. 103. An older and Innger one at tho Hme uporlet. ahnwliig hrsncliea. tcnn left 
baiaor[woMch><ciuiartlie*ev«>n, ■ndllieteriuinil ^UlI balwecn tliem, for ttie uin- 
Uniullon oftbt (Towtli of the miilBlnrl 

FIG. 1M. BnMofitERiorUelLBnMiui 

FIG. ins. 'MoMtrodtyofa PolBto-rli 
the GardaiMn' Chnmicla.) 


So., I 

m Anichoke, ileTcloped 

» represented by obvloiu leaie*. (From 


be called a Monomerous tuber, nomoli in Nclumbium luteum 
(Fig. 108') , where it ooaaists of a eingle thickened mttniode of 

n a(|im1ie niimer. whii-h ia nernpdingly quite destitute of scales 
r buds. TliL- growtli prweediug fVom thia sin)|)le tuber is 
uecesBorily from a bud of 
the node at ils ajM-x, vhencG 
also a cluster of roota is 
pixKiuced. or a somewhat 
similar nature are the con- 
catenate tuln-rs of Apios 
tiilxjrosa (sei'Pi-al of whicli 
10*" are strung as it were upon 

a long Hlirorm axis), llie tulwrs not unfVcqucntiy being mo- 
nomprous. although the larger ones are not so, 

1 1 7'. Taborclect, as they may be tei-med, are of a mixed or 
ambiguous eliarm^ter between tubers and tulierous roots. A good 
example of tlio latter is offonleil by Dahlia-roots. (Kig. 68.) 
They j-iekl their nourishing substance to growing buds on the 
Bt«m almve, but do not themselves normally [irodiice even 


aclveDtitioos boda. Sweet potatoes (55), although equally 
roots, do produce adventitious buds, especially troia near the 
upper end. The somewhat similar tuberdei or tumefied roots 
of certain Orchises ami other plants of the same tribe,' definite 
in number and shape, and sometimes imitating a conn, are 
chained with a bud at the upper end, near their origin. Ap- 
parently, the origin is a bud from the base of the parent 
stem, which bud directly forms a tumcfietl sliort root from its 

118. A Com (Cormat) is to be compared on the one band 
with a short rootstock or tuber, on Ihe other with a bulb. It 
is a siihtcrraiiean fleshy stem, of rounded or depressed figure 
and solid texture. Some of its buds grow into new eorms, and 
these, upon the death of or separation from the parent, become 
new individuals: some deielop abo\e gmimd 
the vegetation and the blossoms nf the season 
A good type of conn is that of (^clamen 
(Fig. 100), in which the >eri base of the 
seedling stem grows flesh} and v- idcns tVum 
year to year, but hardly at all lengtlicns and 
so becomes ftir broader than liigli or de 
pressed. As the main bulk Itelongs to the «» 

first intemode, or caulitlo tlie buds from which tlie \ early 
growths of leaves and flower stalks spnng are at the centre of 
the summit or Hp[>er surface, tlie roots 
from the lower, and the sides seldom pro- 
duce any buds. The corm of Indian , XfifeA 
Turnip (Arisiema triphyllum. Fig. 110) ^ *s3^^i 
is somewhat similar, but it sends up a 
single stout stem, and the roots spring ' 
from around the base of this. These are 
completely naked oorms. 

119. But in Crocus (Fig. 111. 112). Colchicnm (Fiji- 117), 
Gladiolus, and the like, the sheathing hnst'S of one or two leaves 
enclose the corm with a membranous-sealy coat, giving it exactly 
the apix-arance externally of a coated bulb. Sucli have been not 
inappropriately named Molid bulb*. In common parlance, tliey 
will doubtless continue to be calle<l bulbs, and even in iK>|>ii]ar 

' Not, howe»er, tuch •* thoee of ApLwtrum. Tipularin, etc., wliich «re 
genuioe corn» or tubcre. 

: Innuch, Beilr. Biol. & Morphol. Orchid. 1863, fide Duchartr?, El^m. 
Bot. 274 

botanical descriptions. In fact, while they dilTer from nnkect 
1 liaviDg some investment, tbej- ditfer fVom tnte Imlbs 
ouly ill llie greater size of the solid axis and the fewness of 
the investing auales; 
the stem or solid body 
making the greater i>art 
of the conn, but a vorj' 
small part of a proj^r 
Imlb. There are, more- 
over, all grailatioiis bc- 
ii) tween the two. 

120. A Balb, as compared with a conn, may be said to bo an 
exceedingly abbreviated stem, reduced to a flat plate, from the 
lower face of which roots are produced, from the upper face, 
leaves in the form of scales: these scales being either reduced 
ond thicke[ie<I lea\'es or tlie tliickcncd bases of ordinary leaves. 
Compared with buds (73), it is a very fleshy bud, usually large 
and subterranean, tlie axis of which never elongates. It is a 

provision for fiiture growth, the stored nourishment of which is 
dcposit«<l in the leaves, or the homologues of leaves, Instead of 
in the stem. 

, the fliw tliln mTelnidng ivBlni remoTRl, thnvlnjt Itieir 
I miurk the iMda. Um *brt*tllnl tfttte of llie IhM jrnr'i r. 
•ir1<>T>)>>Klntni>ew<uiMMiTu1'>Mi*ntonialur&oe. 113. 
xirm. Willi ■ Mrmlnikl mhI nn« iMarnl biul. 
. S'wtlnn nrktunfcnUil t>Bll> nf Mia Onion. 

. Vcrtlcnl Hiptinn nf (li« hnlb of tlo Tollp, iliuvlne i'> tHm or tinmliu] 
III two iull>ar)i litiiti iK b>. 

n of ■ Uarlli'. •Ith a crop nf yoaiic bolbL 

ttksl wriliHi urUiii Fnrm nf • OnoaT a. new Innti. 

tint •nrllnn nf Ilw conn nf Cukliliiini lA). wILh Ibe wtthfrad aam of 

, anit ilK furmlm nna (t| for ih« cn«oln< jrar. 


121. A. TiuiMted or Coated Bnlb (Fig. 113-115) is one in 
which the scales are broad and completely enwrapping, forming 
concentric coatings. These are thickish when fresh, but thin 
when exhausted and dry, as in the Onion, Garlic, and Tulip. 

122. A. SuIt Bnlb 
has the bulb-scales 
row, thick,and small, 
imbricated, but not 
severally enwrapping , 
each other. That of [ 
the Lily is the most ' 
familiar and cbar-^ 
acteristic example. -^ 
(Fig. 118, 119.) "" u» 

123. Balbleta are small aerial bulbs, or buds with fleshy scales, 
which arise in tlie axils of the leaves of several plants, such as 
the common Lilium bulbifcrum and 
L. tigrinum, the Tiger Lilies of the ' 
gardens (Fig. 120). Here Ihey ap- 
pear during the summer as axillary 
buds: they are at length detached, 
and falling to the ground strike root, 
and grow as independent plants. In 
the common Onion, and in many other 
species of Allium, similar biilblcts 
take the place of flower-buds in the 
nmbel. IJuIblets plainly show the identity of bulbs with buds. 

124. All these extraordinarj', no loss than the ordinary, forms 
of the stem, grow and branch, or multiply, by the development 
of terminal and axillarj' buds. This is jierfectly evident in the 
rhizoma and tuber, and is equally the case in the corm and bulb. 
The stem of the bulb is usually r«luced to a mere plate (Fig. 
114 a), which produces roots from its lower surface, and leaves 
or scales from the upjwr. Besides the terminal bud (c), which 
usually forms the flower-stem, lateral buds (i, 4) are produced 
in the axils of the leaves or scales. One or more of these may 
develop as flowering stems the next season, and thus the same 
bulb survive and blossom from year to year; or these axillary 
buds may themselves be<tome bul1>s, feeding on the parent bulb, 
which in this way is often consumed by its own olTspring, as in 

FIG. Ilg. nciilrbn1bofCanadnlJlT.LIIIuiiiri»iulenK,nneraowerln|. 110. Var- 
UcaI HcUm of Kime. (hnwlng IHO nvvr yiiuHK ImltH- wtlLln. 

FIQ. m. Bnlblalaliittaeullof tboctiulliiclEavHorXlgerLllT. 



the Garlic (Fig. Ilo) ; or, Itnallr separatini; from the Im 
paivnt, just as the hulbleta of the Tiger h\\y fall from tb<' stem, 
they may rorm so many intlepemlent iiulividnals. .So the eonn 
of tlie Crocu3 (Fig. Ill, 112) protliiuoa one or more new ones, 
wliicli feed upon and cxhauat it, and take its plaec ; and the 
next acaaon tJic slirivclleti remains of the old corm may be funiKl 
miderneatli the new. The eonn of Colchicum (Fig. 117) [iro- 
duees a ncn bad on one side at the base, ami iti consumed by it 
in tlie course of the season ; the new one, after Boweriitg by it« 
tenninai bnd. is in turn consumed by its own offspring; ; and 
ao on. The fignre representa at one view, a, the dead and 
shrivelled corm of the year preceding ; ti. tliat of tlio pnisent 
season (in a vertical section) ; anil, e, the nascent bud for tlie 
growtli of the ensuing year. 

125. Condensed Stems, liomologous with corms, tubers, &c., 
and similar in mode of growth, but above grouud, and multiply- 
ing in the same ways, are not uncommon. The Cactus family is 
mainly cuui|M)sed of such forms, of flat- or round-jointed Prickly 
Pears (0[)untia). flut«d or angled eolunnis (Ccreus). and gloli- 
ular Melon-Cactus, Mamillai'in, and Echinocaetus. Tlio latter 
tj-pes, which completely imitate coiins, are the most consolidated 
forms of vegetation. \Vliile ordinary plants are constnietwl on 
the plan of great expansion of surface, these present the least 
possible amount of surface in proportinn to their bulk, their 
l>crmanent siiherieal figure being that which exposes the smallest 
portion of their substance to the air. .Such plants are evidently 
adapte<1 to very dry regiont; and in such only are they naturally 
found. Similarly, bul)>oiis and corm-bcaring plants, anil the 
like, ai-e a form tif vegetation which in the growing season may 
in the foliage exi>and a lar|{o surface to tho air anil ligiil, while 
during the jierioil of rest the Ii\-ing vegetable is redufcd to a 
globular or other form of the least suifaeo ; and tliis is prot«ot«l 
by its outer coats of dcoal ami dry scales, as well as hy its snht«r- 
rsncau situation; — thus exhibiting another and very similar 
atlaptatiiiu to a season of drought. And such plants mainly 
l>elong to countries (such as Soiithern Africa, and the interior 
of Oregon and California) which have a long hot season, during 
which little or no rain falls, when, their stalks and foliage above 
and their roots l)cneatli being early cut otf liy drought, the plants 
rest set^urcly in the conn-like forms to which they ai* Rvluccd, 
and retain their moisture with great Icuaeity until the rainy season 
returns. Then they shoot forth leaves uud flowers with wonderl'til 
ra|iidity, and what was perhaps a desert of arid sand beeomoa 
green with foliage and gny with blossoms, almost in a tlay. 



126. Stems seiring the purpose of foliage, Phyllocladia. Most 
of these condensed and permanent stems are illustrations of 
this, their green rind doing duty for leaves, which 
are either absent, or transient, or reduced to 
spines or other organs not effective as foliage. 
In the flat and broad-jointed species of Opuntia, 
and still more in Phyllocactus and Epiphyllum, the $^ 
forms assumed give a considerable surface of green 
rind, which well answers the purpose of leaves. 
Flattened stems or branches of the same sort and 
economy not seldom occur in other than flesh}' or 
succulent plants (such as the Cactuses) ; some- 
times accompanied by a certain number of real 
foliage-leaves, but these more or less transient, 
as in Bossiiea and Carmichselia among L^u- 
minous shrubs, and Muhlenbeckia platyclada, now 
in common cultivation (Fig. 121) ; sometimes 
with all the leaves reduced to small and function- 
less scales, as in the Xylophylla («. e. wooden- 
leaved) section of Phyllanthus, and in Phyllo- 
cladus (New Zealand and Tasmaniau trees of 
the Yew family) . In all these, the cauline nature 
is manifest by the continuous or proliferous 
growth, by the marked nodes and intemodes, 
and often by the bearing of flowers. m 

127. Special foliiform Branches, Cladophylla, are more decep- 
tive, and commonly pass for leaves. They are well shown by 
Ruscus, M}Tsiphyllum, and Asparagus, — all of one family. In 
all these the proper phylla (12) of the shoot are little scales, one 
to each node, and functionless : from the axil of each there 
immediatel}' develops a boiiy in almost ever}' respect similar to 
the blade of a leaf, subsen-ient to the same purpose, and equally 
definite in growth. That is, it has a definite form for each 
species, and is not continued by any ulterior growth from sides 
or apex. It is usually an axillary branch, of one intemode, all 
further development being permanently arrested. In M^-rsiphyl- 
lum, this false leaf is soft and pointless : in Ruscus (called 
Butcher's Broom in I]ngland, Fig. 123), it l)ecome8 firm, hard, 
and spinj'-tipped : in both the cauline nature is evinced by the 
axillar}' origin ; also by vertical instead of horizontal expansion, 
though this peculiarit}' is shared with most leaf-stalks when they 

FIO. 121. Foliiform branch of Mahlenbeckla platyclada, growing from the apex, 
bearii^ a ■mall and transient leaf at some nodes, also a flower or two. 



assume the character and fuDCtion of the blade. (217.) The 
dadopht/U of RuscQS reveals ite character farther by Ix'^ring a 
flowLT on the middle of one face, 
in the axil of a seale-kaf, bo that 
this amy consist of two inter- 
nodes ; or else the llower-etalk 
may belong to an accessory axil- 
lary bud, bear the scale-leaf close 
to the flower, and he cungenitally 
adnate up to this point with the 
axis of the cladophyll. In 
MjTsiphjUiim (a South AfHean 
climber, common in greenhouse 
cultivation under the erroneous 
name of Smilax, Fig. Vii), the 
cladophyll exactly counterfeits a leaf in form and texture, as well 
OS in function, and it never l>e&rs either Gcule-lcaf or blossom; but 
the flowers are on slender stalks from accessory buds out of the 
same axil. The true leaves are represented by thin and minute 
scales which may escape notice. 

128. To all such branches which imitate leaves, Bischoff has 
given the name Pbtllocladia, sing. FirrLLOCLADniM. To lliose 
definitely restricted to one intemode, and which so closely 
counterfeit leaves, Kunth gave the name of Cladooia, sing. 
C1.AD0DIUH. The best common name for all productions which 
imitate leaves would have been that of phyllodium 
(meaning simply a leaf-like body) ; but that term 
was first applied and is restricted to the case 
of a petiole imitetiog the blade of a leaf. The 
name PhyUodadium (meaning a leaf-like branch) 
may properly be retained for the whole series of 
leaf-like bodies here described. But for those of 
the preceding paragraph, which are so peculiarly 
leaf-like, Kimth's name of Cladodium (1. e. a 
branch-like body) is false in meaning, and may 
be replaced by that of Cladopiitlh:m (t, t. leaf- 
branch), or in shorter English Cladopiiyt-l. 

129. FroDdose Stems. Finally, in some few 
phfenogamous plants, the whole vegetation is re- 
duced to a simjtle leaf-like expansion, as in Duckweed (Lemna) , 

riQ. Ill Mrnlpfayllam. with eltdirpbylli Hrving tar fotiacs; tin tr 
•laUngarnilnals nml twy IncoMplciloin waits dnWrndltiB tlio foft " 

Flo. 113. AMngleolulopbjIlurRDMnBicalWaniDUta " ' 
•noUiDTtcste-leufnii the middle of tu b», uid flDircnIiil 

FIQ. XU. Lsnilia mlnol, a common Duakvued, wbuU pi 


Fig. 124. Here is no differentiation whatever into stem and 
foliage ; bat the expanded floating body which serves for both 
mnat be counted aa stem developed horizontally into a flat plate, 
for it produces a root fVom the under surface and a flower trom 
the edge. This simplification is common in some orders of 
Cryptogamons plants ; and such a body, which answers both for 
stem and foliage, is termed a Frono, from the Latin from, which 
means either leaf or leat^' bough. In some species of Lemna 
the frond is thickened or plano-convex : in WolfBa, the simplest 
and smallest of pbsenogamous plants, it is a globular green mass, 
seldom much laigcr than the head of a pin, wholly destitute of 
root, propagated by proliferous budding from one side, and trom 
within the top producing a flower or pair of flowers. 

§ 8. Iktebnal Strcctche. 

130. The investigation of the intimate stnictare of the stem, 
as of the other oigans, belongs to vegetable anatomy or histolc^y 
(treated in Part II.) ; but tie general outlines of structure, so 
far as is requisite to the explanation of what is ^-isible to the 
naked eye, should be here explained. 

131. The stems of phienogamous plants anatomically consist 
of two general elements, the cellular and the woody ; the former 
excmplifie<l in the commoner stems by the pith and outer bark, the 
Utter by the wood. Both are equally composed of cells, or origi- 

nate as such ; but those which form the woody system of the stem 
mainly ondeigo, at a very early period, transformation into tubes, 
some of which are of such small calibre that their common name 
of fibres is not inappropriate ; others, of larger size or ampler 
calibre, take the name of ducts or vessela. The latter arc almost 

no. IM*. Aini^Ulei1al1eeattpnrtlonaftlie<]ow«T-st>lkof KIchkrdlBjRtblopka 
|Uh (o-ckHsd OUb Lily), twurarM with aoms loD(J tudlnal view : mUnlT pan<Dchyrat, 
Q» caU> bnJlt Dp to u to Imva cainiarmtlrelr Iuys Tmcudn (InUniellalu' apuM or 


always associat(>d ^ith the wood-cells, so that they are in a general 
way taken together as constituting the wood, or woody tissue, and 
as forming what is more definitely termed fibro- vascular tissue or, 
when distinguishable into threads, fibro-vaacular bundles. These 
run lengthwise through the stem, sometimes as such separate 
llireads, sometimes confluent into a compact structure. The 
soi^r or at least the non-fibrous portions, formed of comparatively 

short and commonly thin-walled cells, form cellular tissue. Its 
onliuary fonn (of rounilisb, cubical, or polyhedral and thin-walled 
cells) is called imrenchiTna. This abounds in herbaceons stems or 
herbaceous paria : in trees and shrubs, woody tissue largely pre- 
vails ! in most herbs, it forms a notable portion ; in some (esi)ecially 

FIO. 123. FlbrD-TDKalikr slemenU. a. Bait«11s(1(Hig wo«I-«lli)ar flbrona buk 
Of UnJBD or Baw-wODd. b. Soido wood^nllg uitd |bglo«t» dnct. >n<I c. ■ deUcbccl 
wooil-coU of the wood of uttng tme. cqnallir BmBnlBed with a. <l. A iji;lacli«l <KauA- 
n>IlfniD>sbaTln2i>rWblurine,>baBliiiUiep«vu11ardii>k-IIkeinacMngs. «. rortiun 
of Muna (bivlDg. /. PorUoD nf a clotlail itnct It'on tlw Vine, nldenUi nuult np or a 
•rrka of ahon cella. f. pjut d[ a amallat doCI«d duct, (bowing no appMntnne nf Rich 
nimi«MlCI<in. k, L Spiml dudls nr riwdi, of Ilia ordinary klml. J. Spin) ilnsl Of 
nanana. k. Dnnlfhiin Coleir, tbo tliroad within aplral or annular twlow, reHculatod 
almn. and hlj(berpMMln|ilBli> the Ktateiir dotted dncL I. Jiavxtnm ImpaUsna, wltb 
tbe opon (plral psHlDg low rtnga at (bo middle. All magulUed Kiniiwliat equallj. 


in certain aqnatic herbs), it is reduced to a few threads or 
vessels, generally delicate, and sometimes obscure. The ac- 
compan^ing anatomical illustrations (Fig. 124', 125, with their 
explanations) will give a general idea of the nature of the ana- 
tomical elements of the stem. 

132. In the forming state, the whole stem is parcnchj-ma ; but 
an early differentiation takes place, converting certain portions 
into woody or fibro-\ aacular tissue. This is arranged in two 
ways, gi\'iDg rise to two kinds of stem in phicnogamous plants, 
which have been termed the Endogenoiu and the Exogenom,^ 
meaning inside and outside growers. 

133. The two plans of stem arc usually manifested in external 
conformation as well as internal structure, and are correlated 
with important differences in embryo, fohage, and flower.* Palms, 
Lihes, Rushes, and Grasses are examples of the endogenous 
class ; the ordinary trees and shrubs, especially those of cool 
climates, and a large part of the herbs, arc of the e^zogenoua class. 
In an exogenous stem the wood occupies annual eoncentnc lasers 
one of each year's growth the centre is occupied b^ a pith 
composed of parenchyma onlj the circumference bj a separable 
bark ; so that a cross section presents a 
series of rings or circles of wood oi 
the first year one nng surroun ling the j 
pitb and surrounded b} the bark 
end<^cnous stem has the wootl in distinct I 
threads or fibro-vascular bundles travers 
ing the cellular system or parenchj-ma with \ 
little or no obvious order, and presenting 
on the cross-section the divided ends of these bundles in the 
form of dots ; these usually (but not always) diffused over 

1 Tenne introduced by DeCandoUe, following the idcsa ni DcsfonUines, 
and which have played an important part in strucfural and systematic 
botany ever aince DeCandoUe adopted these namcB as those of the two 
primaiy diviiions of phenoganiolu plante, Ezogena and Endogina-. But it 
hat long been aeen that tlie name of the second kind ia not appropriate ; and 
the older and better (though longer) names of Jussieu, Monocotylcdonca and 
Dicotyledonea. are reverted to. Yet the CandoUean names arc still much 
employed, with doe explanation, to designate the two Unda o( atructure of 
the Btem. 

* Yet with some more or tees valid eiccptions, aa when the annual stem 
of Podophylluni and the rhizoma of Nymphsa, among dicotyledonnua plants. 
Imitate the endogenous structure ; or where the pith of an evidently eiogenoua 
■tern, ai in the Piperaces, haa aeattered woody bundles in an cndogenoua 
fashion ; or where monocotyledonous plants have all their woody bundles in 
■ deflnile drcle, aa in Luzula, Croomia, &c. 

no. Ua. BectloaoraimaUPaIm-«lsm, In twodlnctJoDS. 


Ihe wLole section, or when few in number of flomewhat definite 
position or arrange mi.' n I. The ordinary' appearance of such a 
Bt4?m. both on the lon^tucUnal and the eross-scction. is shown 
in Fig. 12C; it may also be examined in the Cane or Rattan, 
the Bamboo, and in the annual stalk of Indian Corn or of 
Asparagus. The appearaiiee of ordinarj- wood U very familiar. 

135. The newer woody bundles of an endogenous stem arc vari- 
ously intermingled with the old. When DcCandolle gave the name, 
it was supposed, from Desfontaines's researebcs, that the older 
bundles occupied, or came at length to occupy, the eircumference 
of the trunk, while only new ones were formed in the centre; 
and that increase in diameter, when it took place at alt, resulted 
Ttoxa the gradual growth and distention of the whole. Henca 
the contrasting name of endogenout, or tnttde grwcinii, and for 
auch plants tLe name of ENtiocEXors Pi.asts, or Bndookks. 
Our actual knowledge of the structure and growtli of these stems, 
as will be seen, cannot be harmonized witli this view in any 
way which gives to the name endogenous an appropriate signifi- 
cation. The name continues a.i a counterpart to the more correct 
one of exogenous, and as a survival of former ideas. 

136. The Endo^nons Structure (so called) of the stem is cor- 
related with a monocotyledonous embryo (89), usually with a 
ternary arrangement in the flower (3'22), and commonly with 
parallel- veined leaios. (173.) Endogens, although they have 
manj' herbaceous and a few somewhat woody representatives 
in cool temperate climes, mostly attain their fiill variety of fea- 
tures and rise to noble arborescent forms under a tn>pical 
Bun. Yet Palms — the arboreous type of the class — do extend 
as far nortli in this country as the coast of North Carolina (the 
natural limit of the Palmetto, Pig. L'26*) ; while in Europe the 
Date and the Chamwrops thrive in the warmest parts of the Enro- 
pean shore of the Mediterranean. The manner of their growth 
gives them a striking appearance ; their tnmks being unbranehed 
cylindrical columns, rising to the height of from thirty to one 
hundreil and fifty feet, and crowned at the summit with a ^mple 
cluster of peculiar foliage. Palms generally grow from the 
terminal bud alone, and jterish if this bud be destroyed ; they 
grow slowly, and bear their foliage in a cluster at the summit of 
the trunk, which consequently forms a simple cylindrical column. 
But in some instances two or more buds develop, and the stem 
branches, rarely and accidentally in ordinarj' species, regularly 
in the Doum Palm of Upper Egjpt, and in the Paulanus, or 
Screw-Pine (Fig. 69), which belongs to a family allied to Pahns : 
in such cases the brunches ore cylindrical. But when lateral 




bods are fVeel; developed (aa in the Afiparagua), or the leaves 
are scattered along the stem or branches by the flill development 
of intemodee (ns in 
the Bamlxm, il:ii;;i.', 
&c.)'they grmlually 
taper upward in the 
manner of moat ex- 
ogenous sterna. 

137. This kind 
of stem comprises 
several subordinate 
types as to int<'rnal 
structure, whtch to 
be well understood 
must be studied his- 
tol<^cally , undertlie 
microscope.' To one 
of these, by do means 
the simplest, belongs 
tiie ordinary iialm- 
stem, the anat^imy 
of which was made 
classical by llohl, 
and has been 
supplemented ^ 
by Na^li. 
In this a large .^ 
part of the a 
bundles, or all 
of the more 

conspicuous kind, starting fVom the base of the leaf to which 
they respectively belong, eune inward more or less strongly 
toward the centre of the stem, and thence gradually outward 
as they descend until they reach the rind, in which the 
attenuated lower extremity mostly terminates. Consequently, the 
bundles from different heights cross in their course, somewhat 

1 For the best and most acccuible memoir on the nibj«ct, of recent dale, 
■ee GnilLaiid, Jlecherchei inr I'Analomie compart et le IMfcloppemeiit des 
TIhiu de la Tige dam ]e» Honocotjlnlnnes, publiahed in Ann. Sci. Kat 
■er. 0, T, 1-1T6, ISTT. Six tjpei of the ilem of MaDocotjIrdoni are here 
recognize hy anatomical character* and mode* of grotrth, one of them 
having four modi&cationi. 

FIQ. llr. Satial ?BlmMto In vaifou ita|<i; alaa tha TBOca aloUbUa or SpaaUi 


as ahown in Fig. 127. It is partly owing to this c 
these fibres vritli tho riod tbat the hitter is not spparable from 
the aU^m. In some- I'ahns, and Lii Grassea, 
h there i» no markiMl distinction bi'tnccn 
I the wood and rind, or no proper rind at 
In others, such as the Palmetto 
I (Fig. 120), there is a mnrkeil rind 
I or false bark, which receives independent 
S fibro- vascular bundles from tlie leaf-stalks, 
A and is Iraverecd by them in [inrallet lines, 
(j In Grass-stems, and others with long iiiter- 
J nodes and closed nodes, the fibro-\-aBcu1ar 
J bundles all run approximately straight 
1 1 and parallel through the int«modcs, but are 
J intricate and anastomosed in the nodes. 
The whole centre of the intcmodes, when 
n'l ■ .1 !,■ it becomes so. ia occupied by a true pith, 

like tti^t ul' un L^iogcn, and in some cases eqnally destitute of 
fibro-vasfular bundl(«, but oltcn with scattered ones, after the 
manner of certain Exogens anomalous in this respect, such as 
Nyctaginaceie and some AraliaceiE. Endogenous atems of 
simpler stniclure, as in herbaceous Liliaccie, Commelynacca;, 
Ac, have a distinct cortical portion (at least in the root-stock 
or portion of stem properly eomparable n-itli palm-trunks and 
the like) ; but this is mostly destitute of iibro-vaacular bundles. 
Most of them liavc two kinds of vascular bundles, one 
of wliich not rarely occupies an exact circle in the line of 
division between the cortical and medullary portion (between 
bark and pith), and the other is wiUiin Ibis circle, either of 
verj- few and scattered bundles, as in Cunvallnria majalis, or 
numerous and scattered, as in Uvuliiria and the leafy elcma of 
Tradescantia Virginica ; or these bundles are few and arranged 
nearly in an inner circle close aronnil the centre. Finally, Luzula 
and Croomia have only one kind of bundles, answering to the 
outer ones of Convallaria ; in other words, the woody system 
forms a simple circle, dii-iding a purely ci-lluUr medulhiry from a 
similar cortical portion, thus closely imitating an herbaceous 
exogenous stem of the same age, 

138. An annual endogenous stem increases in diameter by 
general growth imlil it attains its limit. Ligneous and enduring 
stems inere.ise similarly up to a certain jieriud. Then the rind 


sooner or later ceKsea to diBtend or adapt itself to Rirtber in* 
crease in diameter, and there is oo interior provision for indefinite 
increase in the greater number of woody endt^enous trunks. But 
in Draciena (Dragon-trees) , in the arborescent Yuccas, and the 
like, the zone intermeiUate between the cortical and interior re- 
gion, which is for a time active in many Endogens, here grows 
continuously and indefinitely. Such trunks increase in diameter 
throughout life ; they may attain a very great age (as some 
Dragon-trees have done) ; and they imitate exogenous trunks 
to a considerable extent in mode of growth. 

139. The wootl of an end<^enou8 woody stem is hardest and 
most compact at the circumference ; in palm-atems commonly 
it is largely mixed with parenchyuia or pith at the centre, even 
in old trunks. 

140. The ExogenonB Stmctnre, that of ordinary wood, is char- 
acterized bj' the formation of a distinct zone of wood between a 
centml cellular medullary portion (pith) and an outer chiefly 
cellular portion (bark), traversed by plates fk)m the pith (medul- 
larj- rays) , and by increasing from the outer surface of this zone 
between wood and bark, the increase in enduring stems consist- 
ing of definite concentric annual layers. 

141. Its Beginning, at the earliest growth of the embryo, is in 
the appearance of a few ducts (Fig. 125,/-^), at definite points 
in the common parencliyma of the initial stem (four equidistant 
ones in the Sugar Slapte) ; each is soon surrounded by incipient 

proper wood-cells (Fig. 1 25 , i, c), K^ether forming a fibro-vascolar 
bundle or thread. Additional ones arc intercalated as the second 
and third intemodes develop, and so a column (in cross- 
wction a ring) of wood is produced, always so arranged as to 

Fia. 128. EHagnm of % cmU'Wctlon of ■ tanaiag mhHId; Mem, ahowliii Uis 
niknner In which Uie ronng wood li urnDged In lbs cellnlir lyitem. 

FIQ. 1:9. Tbe ume at ■ Inter [«riuil, tba voodj bonills lacreueil (o u nsnrif to 
flu tbe rlrele. 

FIO. 130. The nme tt Ibe close Df tbe isuan, wbare lbs wood \iu fonncd a com- 
plete drde, Intempted mij bj tbe nednODir nqri, wUdi ndUte from the pith to 
the bark. 


surround a purely cellular central part (the pith), while bui>- 
roundetl by a cellular external rind, the bark, or outer bark. 
The diagmms (Fig, 128-130) rudely sliow somo stages in the 
formation of the zone of wood. The fibro- vascular bundles 
originate in the bases of the leaves, and develop outward in{« the 
forming leaves as well aa downward into the forming stem. 

142. First Tear's Growth. The woml, even in a herbaceous 

or annual stem, at the completion of the first year's growth, 

ui forms a zone or tube, 

I enclosing the pith. Dut 

it is traversed by plates 
(in cross-section lines) 
of parenchyma, or cel- 
lular tissue of the same 
nature as the pith, 
which radiate from 
that to the bark, and 
thus diridc the wood 
into wedges. These 
lines, forming what is 
called the n'lver-ffrain 
in wood, are the Med- 
CLLAJiT Rats. They 
represent tJie cellular 
system of the wood it- 
si'lf, or un transformed 
parenchyma. Beiug 
pressed by the woody 
wedges, their eelb are 
laterally flattened, tn 
some stems, the med- 
" ullary raj's, or many 

of them, are comparatively broad and couspiaious ; in others, 
thin and inconspicuous or irregular. The growth of the woody 
wedges is soon complete, except at the outer portion, next 
the bark : here tliey usually continue to grow llirougli the 
season ; that, is the wood grows externally. The general ana- 

Fta. 131. LontluiiUiu] unl U 
dApynrpnm )< mt Che cIobs of the flnC iti^Vh jETOwth i of t 

FIO. 132. Portion 0ft1ieunie,mi^lfl«l,ahaii1nEt)i 
Ota wwd, wid UaI by tfas Inrk. 

FIO. 133. Man nKgnifled dlni of U: 

a. part of ttia pitb ; b, ti 

I at tlio Son M*pla (Acer 
lulu |<itli, RucroawSed bj 

.b; c.X1iewaod;if,<I. •)< 

Ibrouii bark ; fr. tba nllultr onrelope, 

' s^ilenali ; k, dus of tba mednltar]' 


tomical strocture of a woody exogenous stem of a year old is 
displayed in the Fig. 131-133. Viewing the parts particularly, 
and in order from centre to circumference, there is, — 

1st. The Pith or Medulla ^ consisting entirely of soft and rather 
large thin- walled cells, ^ gorged with sap or other nourishing 
matter during the growing state, becoming light, dry, and empty 
when effete. 

• 2nd. The Layer of Wood^ traversed by the medullary rays. In 
Pines and other Coniferaj, the wood is of uniform structure, being 
wholly composed of a woody tissue with peculiar markings (Fig. 
125, rf, e) : in other wood, ducts of one or more sorts occur ; the 
most conspicuous being what are termed dotted ducts. These 
are so large as to be evident to the naked eye in many ordi- 
nary kinds of wood, especially where they are accumulated in 
the inner portion of the layer, as in the Chestnut and Oak. In 
the Maple, Plane, &c., they are rather equably scattered through 
the annual layer, and are too small to be seen by the naked eye. 
Next the pith, i. e. in the very earliest formed part of the wood, 
some spiral ducts are uniformly found, and this is the only part 
of the exogenous stem in which these ordinarily occur. They 
may be detected by breaking a woody twig in two, after dividing 
the bark and most of the wood by a circular incision, and then 
pulling the ends gently asunder, when their spirally coiled fibres 
are readily drawn out as gossamer threads. As these spiral 
ducts form a circle immediatelj' surrounding the pith, they have 
collectively been termed the medullary sheath, but they hardly 
deser\'e a special name. The vertical section in Fig. 133 divides 
one of the woody wedges, and shows no medullar}' ray ; but there 
is one at the posterior edge of the transverse section. But, in the 
much more diagramatic Fig. 134, the section is made so as to show 
the surface of one of these plates, or medullary rays, passing hori- 
zontally across it, connecting the pith {p) with the bark {b) . 
These medullary rays form the sHver-grain (as it is termed) , which 
is so conspicuous in the Maple, Oak, &c., and which gives the 
glimmering lustre to many kinds of wood when cut in this direc- 
tion. A section made as a tangent to the circumference, and 
therefore perpendicular to the medullary rays, brings their ends 
to view, as in Fig. 135, much as they appear on the surface of a 
piece of wood from which the bark is stripped. They are here 
seen to be composed of parenchyma, and to represent the horizon- 

1 In rare instances, a few fibro-TascuUr threads are found dispersed 
through the pith, presenting a somewhat remarkable anomaly. This 
occurs in Aralia racemosa, and more strikingly in Mirabilis and other 
Nyctaginaceae, and in Piperacese. (163, foot-note.) 


tal Hystem of the wood, or the woof^ into which the vertical woody 
libre. &c., or wnrp, is interwoven. Tlie inspection of a piece of oak 
or maple wood at ouce shows the pertinency of this illnstration. 

3rd. The Bark or rind. This at first consisteil of simple 
parenchyma, like that of the pith, except for the green color 
develojjed in it, the same as that which gives verdancy to foliage. 
This green matter is formed in the ceils of all such parts when 
exjHised to light, consists of green grains of somewhat complex 
chemical coiii|K>sition, has imiwrtant t\mctions to perform in 
assimilation (■'. e. In the conversion of the plant's crude food into 
vegetable matter), and is named Ciilui{Ophii.l, i. e. leaf-green. 
The completed bnrk, when all its |)aits are ajjparent, as espe- 
cially iu most trees and shrubs, is composed of three strula, of 
which the green baik, the roost conspicuous in the young shoot, 
is the middle layer, therefore named tlic MESorm/Eiii. This is 
soon covered, and the green color obscured, by & superficial 
stratum of cells, generally of some shade of ash-color or brown, 
occasionally of brighter tints, which gives to the twigs of trees 
and shrubs the hue characteristic of each species, the Corky 
ENVELOPE or layer, or Epiphi/eum. The latter name denotes its 
e.xternal position ; the former, that it is the laj'er which, when 
much dewloped, forms the cork of Cork-Oak and those corky 
expansions which arc so conspicuous on the twigs of the Sweet 
Gum (Liquidambar) , and on some of our Elms (Ulmus alata 
and racemosa). It also forma the imper-like exfoliating layers 
of Birch-bark. It is composei) of laterally flattened parenchy- 
matous cells, much like those of the Epidermis (Fig, 133, (')• 
which directly overlies it, and forms the skin or surface of the 


134. VBrtlal •wtlw 








■bow ona of the mi'ii 



ely rrera 

tbe pHb 

Pi to 







)iIhIs HiHtclilDg ur< 



ul (ectlon ncron ibe ends 







stem, aod of the whole plant. Lastly, the inner bark, accord- 
ingly named ExDOPnL<EUM, takes the special name of Liber, and 
is the most important portion of the liark in the stems of trees 
and shrubs. Complete and weU-dfveloped liber, like that of 
Linden or Basswood, contains two peeuliar kinds of cells in 

addition to comrton parenchyma. Ixtth of the fibrous or vascular 
class: viz., 1. Ckiurifohm or Sieve-cells, a sort of duets the 
walls of which have open slits, tlirotigti wliich they communicate 
with each other; 2, Bast or Bapt-cem^, the flbre-like cells 
which give to the kinds of inner bark that largely contain them 

FIO. I3B*, Portlnn ofs inin>ven«KCtloTi [(bore), anil acniTtfipaniUngTertlCBlMc- 
tlnnlbelow), iDRgnltlol. reuhliiu rrom the |dItar/>) to the vpUlcrmli (r) ol a Mem 01 
VeennAo. a learnlik /). tbebark; JT. tbawoal; anil C. thscnmbliim-lairar. aifiiuncl 
rt> nrcrred to by ■mall letter* are: p. ■ lurllon of (be pith: 
■nslullarr nj where It rniwlnlo tlie|>itb: four com |>1ete ineil- 
a IraniTenq ivrtlnn, appear In (he upper flgiirc, running from 

In Febrnarr. The r 

ptib tn bark : au. TnoliillarT en 

wtMWIIh nncolllnj eitrsmlty 

dotlad iluct* Intenpersnl In (De noou: cl. camDIam-uijer or i 

wood and Inner bark : l-t. llh«r or inner bark, [be Inner portion of 


ij ti», 

' ontei cellular baik; 



their strength and toughness. They are like nood-eellfl except 
in their greater length and flexibility-, and in the thickness of 
their wallH, which greatly exceeds the calibre. This is the 
malerial which gives to the bail or inner bark of Baaswood, &o., 
the strength and pliability that adapts it for cordage and for 
making mats : it is the material of hnen, and the like textile 
fibres. (For a vitiv of the whole composition and structure of a 
woody stem at the close of the Brst year's growth, end immedi- 
ately before that of the second year begins, see Fig. 135*.) 

14.3. Annnal Increase In Diameter. An herbaceous stem does 
not essentially differ from a woody one of the same ago, 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 proi-ision for continuing its 
growth from year to year. As the layer of wood continues to 
incren^ 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 eel I -multi plication is merely sus- 
pended, so there is always a zone of delicate j^oung cells in- 
terposed between the wood and the bark. This is called the 
Cambivu, or, better, the Cam uinn- later. It is charged with 
organizabic matter, which is particularly abundant and mucila- 
ginous in spring when growth rerommences. This rancilaginons 
matter was named Camlnum by the elder botanists : they sup- 
posed — as is still popularly thought — that the bark, then so 
readily separable, really separated from the wood in spring, that a 
quantity of rich mucilaginous sap was poured ont between them 
and became oi^anizcd into a tissue, the inner part becoming new 
woo<l, tlie outer, new bark. But deheate slices show thot there 
is then no more interruption of the wood and inner hark 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 effeeted by the rending of a 
delicate forming tissue. And if some of this np|>arent mucihige 
be scraped off from tlie siirfhce 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 rae<lullnrj- rays here and there in- 
terspersed. The inner jwrtion of the cambium-layer is therefore 
nascent wood, and the outer is nascent bark. As the ceils of 
this layer multiply, the greater number lengthen vertically into 
woody tissue : some are transformed into ducts ; and others, 
remaining as parenehj'ma, continue the medullary raj-s or com- 
mence new ones. In this way, a second layer of wood is formed 
the second season over the whole surface of Uie former layer 


between it and the bark ; and this is oontinuous with the woody 
layer of the new roots below and of the leafy shoots of the sea- 
son above. Each succeeding year another layer 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 occupjdng the circumference. Each 
layer being the product of a single year's growth, the age of an 
exogenous tree may, in general, be correctly ascertained by 
counting the rings in a cross-section of the trunk. ^ 

144. Demarcation of the Annaal 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 layer, where their 
open mouths on the cross-section are conspicuous to the naked 
«ye, making a strong contrast between the inner porous and the 
exterior solid part of the successive layers. In maple and beech 
wood, however, the ducts are smaller, and are dispersed through- 
out the whole breadth of the layer ; and in coniferous wood, viz. 
that of Pine, Cypress, &c., there are no ducts at all, but only a 
uniform woody tissue of a peculiar sort. In all these, the de- 
marcation between two layers is owing to the greater fineness of 
the wood-cells formed 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 arc most distinct in trees of temperate climates like 
onrs, where there is a prolonged period of total repo^, 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 Sipho, 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- 
I>osed, 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. 


vigorons vernal growth, with much larger cells, thuB narking 
an abrupt transttion Trom one layer to the next. Besides being 
finer, the later wood-^eUa of the aeason are commonlj- flattened 
antero-poBteriorlj", probably by growing under grcatt^ pressure. 

145. Each layer of wood, once formed, remains essentially 
unchanged in position and dimensions. But, in tmnks of con- 
siderable age, the older layers undergo more or less change in 
color, density, perviousness to moisture, &c. 

146. Sap-wood (Al.nuBSUM). In the plantlct and in the 
developing bud, the sap ascends through the whole tissue, of 
whatever sort : at flrst through the purenehj'ma, for there is then 
no other tissue ; and the transmission is condnucd through it, 
e8])ecially through its central portion, or the pith, in tlie growing 
a|>ex of the stem throughout. But, iu the older ]>arts below, the 
pith, soon drained of sap, becomes tilled with air in its place, 
and thenceforth it bears no part in the plant's nonrishinent. As 
soon as wood-cells and ducts are formed, they take an active 
part in the conveyance of eap. for which their tubular and ca- 
pillary character is especially adapted. But, the duets iu older 
parts, except when goi^ed 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 tliem, the older ones are each year removed a 
step farther from the region of growth ; or rather the growing 
stratum, which connects the iVesh rootlets that imbibe with the 
foliage that elaborates the sap, is each year removed farther from 
them. The latter, therefore, ader 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 nails and smaller calibre than those of the newer. 
Thus arises a distinction — sometimes obscurely marked, some- 
times abrupt and conspicuous — into sap-xcood and heart-wood. 
The former is llio popular name given to the outer and newer 
layers of soUer, mure oi>en, and bibulous wood. The early physi- 
ologists nametl it nibvm«ni from its white or pale color. Being 
more or less snppy, or containing soluble oi^anic matter, and 
readily imbibing moisture, this part of the wood is liable to decay, 
and it is therefore discarded from timber nsetl for construel ion, 

147. Heart-wood (or Duramen, so called from its greater hard- 
ness or diu-abilily) is the older and mature portion of the wood. 


In all trees which have the distinction between the sap-wood and 
heart- wood well marked, the latter acquires a deeper color, and 
that ijeculiar to the species, such as the dark brown of the Black 
AValnut, the blacker color of the Ebon}', the purplish-red of Red 
Cedar, and the bright 3ellow 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 
deca}', as in the AVhite Pine, Poplar, Tulip-tree, &c. The 
heart-wood is no longer in any sense a living part : it may perish, 
as it frequentl}' does, without affecting the life or health of the 

148. The Growth and Dnration of the Bark^ also the differences 
in structure, are much more various than of the wood. Moreover, 
the bark is necessaril}' 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 decav 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 disapjjear by slow decay. So that the bark 
of old trunks l^ears 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 
readil}' 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 3'car ; the opaque corky layer soon excludes 
it from the light ; and it gradually perishes, never to be renewed. 
The corky layer usuall}' increases for a few years only, by the 
formation of new tabular cells : occasionall}' it takes a remarkable 
development, forming the substance called Cork^ as in the Cork 
Oak, and the thin and parchment-like layers of the AVhite 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-la\-er applied to its inner surface. Sometimes this 
growth is plainly distinguishable into laj'crs, 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 diflTerent 
plants, especially in trees and shrubs. That of Basswood or 



^^— depc 

Linden, and of other plants with a BuniUr fibrous bark, may be 
taken as best represeDtiug the liber. Hero it consists of alter- 
nate strata of fibrous bast, and of the pecnliar Uber-cells called 
sievc-ceUs, in which nourisbiug matter is especially contained 
and elaborated. ^Vhilc the latter, or their equivalents, ouenr 
and iilay an important part in all inner bark, the bast^cells are 
altogether wanting in tlie bark of some plants, and are not pro- 
duet^l utter the first year In many others. The latter is the case 
in Neguudo, where abundant bast-cells, like those of Basswood, 
compose the exterior portion of the first year's liter, but none 
whatever are formed in the subsequent layers. In Beeches and 
Birches, also, a few bast-cells are produced the first year, but 
none afterwards. In Maples, a few are formed in succeeding 
years. In the I'ear, bast-cells are annually formed, but in very 
small quantity, compared with the pai-enchjTnatous part of the 
liber. In Pines, at least in White Pines, the luirk 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 
Uber-celle in their structure and markings. 

151. The bark on old stems is constantly decaying or falling 
away from the surface, without any injury to the tree ; just as 
the heart-wood within may equally decay without harm, cxcejtt 
by mechanically impairing the strength of the trunk. There arc 
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 many years' growth. 
although only the innermost layers are alive : in others, it scales 
off much earlier. On the stems of the common Honeysuckle, of 
the Nine- Bark (Spiroia opulifolJa), and of Grape-\ines (except 
Vitia vulpioa), the Uber lives only one season, and is detached 
the following year, hanging loose in pnjx.Ty layers in the former 
species, and in fibrous shreds in the latter. 

152. While the newer layers of the wood abound in crude sap, 
which they convey to the leaves, those of the inner bark abound 
in elalmraled sap, which they receive from the leaves and convey 
to ttic cambiuiu-layer or zone of growtli. The projier juices and 
peculiar products of plants are accordingly found in the foliage 
and the Imrk. especially in the latter. In the hark, therefore 
(either of the stem or of tlie root) , medicinal and other principles 
arc usually lo 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 origiQat« in the former are 
deposited in the wood. 


153. The LiTing Parts of a tree or shmb, of the exogenous 
kind, are ob\iously 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- 
laj'er, which, annually renewed, maintain a living communication 
between the rootlets on the one hand and the buds and foliage 
on the other, however distant the}' at length may be. These are 
all that is concerned in the life and growth of the tree ; and these 
are annuall}' renewed. The branches of each 3'ear's growth are, 
therefore, kept in fresh communication, by means of the newer 
la3'ers 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 conve3'ed 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 parti}' in the formation of a new layer of wood, reaching 
from the highest leaves to the remotest rootlets. 

154. LongeTitj of trees. As the exogenous tree, therefore, 
annually renews its buds and leaves, its wood, bark, and roots, 
— ever}' 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 years, and some are possibly 
older. ^ Equally long may sumvc 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 Teneriflfe, now destroyed 
by hurricanes and other accidents, had probably reached the age 
of more than two thousand vears. 

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 fVom casualties or attacks to 

* 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 V<^gc'tale ; " more recently, by Alphonse IX'Can- 
dolle in the " Biblioth^que Universelle ; " and in this country by myself in the 
*' North American Review," for July, 1844. For an account of the huge Red- 
woods (Sequoias) of California, see Whitney's Yosemite Book. 

H4 Moeraoi/iGT of stems. 

r-r*!i^t, A tr»* lik* tk^r BaxsvaD (->>. Fsg. 71 1. wh;..-h r.-j aerlil 
rtffAfi Profit ii*w^ to fono iwrw trunk.<^ for tiMr •afjfj»>rt aetI •astenaa*.-!* 
oftUf: *i\>r*iikfViTi'jL hr^u':iifr^, aiiyl ihwi ever a^iv&iK^^s into ikrw •«j:1. 
Ki2M a tnjiv uA*-f:uAjH exUtffWje : bat, tb«rD. it fj<<i>3ie$ a f«>xv>t. 
or j.«i 1// Ur \lkf:n<if\ to a f-r^k/nv prr^pagate^l aii^l in-lrfiiiitcly in- 
ritAv^l ffV -iiH.-krrrsf. ofTs^it. or rnber .saltern nc-an «h«>>t.s. >•.■ 
thf: i^wMion of t^if; f^nfuhiT cr>ntinimtioo of the in^litifiaal plant 
\f4:i:0fttnrn twrr^iA in that of ^:^^t in nation of the rartr. — at Iea>t of 
a \Av\'\fTo\fA^HUA nu:i(T. — the aaswer to whieh is whollv in the 
riornain of c-^inj^r^rtufff.' IIowe%'er this may be, it is e\i«lent that 
a vfrjfr'taMi; of the higher jfra^Ie is not justly to \fe cjmitart*! with 
an aniinal of higher gra/le: t^iat in<li\idaality is incompk'tely 
rt:B\mA in the vfrgetable kingdom :^ that rather 

] hit. Iht. Fliuit in a CmapcMiite Beim^, or Comaiuitj, lasting, in 
the eaxi; of a tnf^r, thniugh an indeGnite ami often immense num- 
\h'T of f^enerationjf. Tlwrsf; are successively prodace<L enjoy a 
ti;nn of exiMt^'urrc*. ami |ierish in their turn. Life passes onwanl 
Cf^ntinually from the older U> the newer |>art8. and death follows, 
with ef|ual Hti'p, at a narrow inter\'al. No |Xirtion of the tree is 
now living that was alive a few 3'ears ago ; the leaves die annu- 
ally tkw\ are cant off, while the intemorles or joints of stem that 
Ixmj them, an to their w^kxI at least, buried deep in the trunk 
under the w^kkI of succeerling generations, are converted into 
lifehrHH heart- wcxxl, or iiercliance decayed, and the bark that 
lNflong(?fl Xa) thern is thrown off from the surface. It is the aggre- 
gat4% the blend(*d mass alone, that long sunives. I'lants of 
Mingle cells, and of a deflnite form, alone exhibit complete indi- 
viduality ; and their exisUfUce is extremely ]»rief. The more 
(romplex vegetabh; of a higher grade is not to lie compared with 
the animal of the highest organization, where the offspring always 
Hcparates from the parent, and the individual is simple and mdi- 
visibh;. Ihit it is truly similar to the branching or arl>orescent 
coral, or to other comixMind animals of the lowest grade, where 
HUccrcHMive generations, though capable of living inde[)endently 
and HometimeH st^parating H])ontaneously, yet arc usually devel- 
oiM*<l in coiniection, blende<l in a general l)ody, and nourished 
more or less in common. Tliiis, the coral structure is built up 
l)y the combined lalnirs of a vast number of individuals, — by 
the HUCccHsive lalMjrs of numy generations. The surface or the 
rc»c<»nt HluH)ts only are alive ; lH*neath are only the dead nnnains 

1 Set' UftrwiiiianA. xli. :i:U^{r>r>. 

> Ah, iMTliiipN, wan flrKt explicitly Htated by Kngelmann, in his innugiirnl 
i»Kiiny, IK' Aiitliolyai l*ru<]n>tuuii, Introduction, § 4. 


of ancestral generations. As in a genealogical tree, only the 
later ramifications are among the li\ing. The tree ditFers from 
the coral stnicture in that, as it ordinarily imbibes its nourish- 
ment mainly from the soil through its roots, it makes a downward 
growth also, and. bv constant renewal of fresh tissues, maintains 
the communication between the two growing extix»mities, the 
buds and the rootlets. Otherwise, the analogy of the two, as to 
individuality, is well-nigh complete. 

Section IV. Of Leaves. 
§ 1. TiiEiK Nature and Office. 

137. Leaf (Lat. FoUum^ in Greek form Phylhun)^ as a botani- 
cal term, has on the one hand a comprehensive, on the other a 
restricteil 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 anv, 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 onlinar}' 
leaf, may naturally In? spoken of as the whole. Petiole and 
stipules are indeed subsidiary when present, and are not rarely 
wanting. Yet sometimes they usurj) the whole function of foli- 
age, and sometimes there is no such distinction of parts. 

15l>. Physiologically, leaves are grcH?n expansions borne b}' the 
stem, outsprt^ad in the air and light, in which assimilation (3) 
and the processes connected with it are carried on. Vegetable 
assimilation, — the most essential function of plants, being the 
conversion of inoi*ganic into organic matter, — takes place in 
all onlinary vegetation only in green parts, and in these when 
ex[X)sed to the light of the sun. And foliage is an adaptation 
for largely incrcasing the green surface. But stems, when green, 
take part in this oflk'c in i)roiK)rtion to the amount of surface, 
sometimes monoix)lize it, and in various cases increase their 
means of doing so by assuming leaf-Hke forms. (120-129.) 
Leaves, especially in such cases, may lose this function, appear 
only as useless vestiges, or may be subser\ient to various wholly 


different uaea. Form and flmclion. therefore, are not sure indi- 
catiuQs of the true nature of organs. 

ICO. Mor|)hologieull,v, and iu the most comprelicnsive sense, 
leavee are B[>eL-ial lateral outgrowths n'om the stem, definitely 
and symmetrically arranged uimu it ; in ordiiiaiy vegetation and 
in tlie most general form constituting the assimilating apparatus 
(or foliage), but also occurring in other fonns and subserving 
various uses. Sometimes these uses arc combined with or eut>- 
sidiarj- to the general function of foliage ; sometimes the leaf is 
adapte<l to special uses only. So the botanist — ree<^nizing the 
essential identity of organs, whatever their form, which apjicar 
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 Uly-bull) or the coat of an onion, the scale of a winter hud. and 
the petal of a blossom. Therefore, while expanded green leaves 
(which may be tautologically termed /o/tuye-Zeai-eg) are taken as 
the proper tj-pe, the common name of leaves, in the lack of any 
avnilable generic word, is in morphol<^cal language exteuflert to 
these special formq whenever it becomes netdful to express their 
l^ylline or foliar nature. 

IGl. In the morphological view, all the plant's oi^ns except- 
ing roots (and excepting mere superficial productions, such as 
hairs, prickles. Ac), belong either lo stem or to leaves, arc 
either cautiiie or phylline in nature. To the latter belong all the 
primary outgrowths from nodes, all lateral proiluctions which 
are not axillary.' Whatever is pi-oduced in the axil of a kaf is 
cauline, and when developed is a branch. 

lf»2. The Duration of Leaves is transient, compared with that 
of the stem. They may bo fugaciow, when they fall off soon 
after their appearani-e ; deciduoui, when they last only for a 
single season : and perii'itrnt, when thej' remain through the cold 
season, or other interval during wliich vegetation is interniptod, 
and until after the appearance of uew leaves, so that the stem is 
never leafless, as in Ertrgreen*. In many evergreens, the leaves 
have only an annual duration ; the old leaves tUlting soon an«r 
those of the ensuing season are expanded, or, if they remain 
longer, ceasing to bear any active part in the economy oC^the 
vegetable, and soon losing their vitality altf^ether. In I*ines 
and Firs, however, although there is an annual fall of leaves 
either in autumn or spring, yet these were the produce of some 

' There are cb»c» in vrhich lliis rule is of difflt-ult upplicaiion, or )■ •eem- 
idIeI}- vfulaled, lamttlliHi hy the iiii|ii>n'Minii at the BUbleniling leaf, as in the 
inflorMcenee of CruvifvriE, rarely in oiIkt way*, to be explaioed in the 
proper plai.'M. 


season earlier than the last ; and the branches are continuall}' 
clothed with the foliage of ftx)m 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 ijeriod, and they all perish in autumn ncarh* 

163. Leaves soon complete their growth, and have no power 
of further increase. Being organs for transpiration, a ver}' large 
part of the water imbibed by the roots is given out b}- 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 diu*ation, at least in 
their active condition. 

164. Defoliation. The leaves of most Dicotyledons and some 
Monocot3*lcdons separate from the stem and fall by means of an 
articulation at the junction with the stem, which begins to form 
early 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, snch 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 mostlj' takes place at 
the approach of winter. In warmer climates ha\ing only winter 
rain, this occurs in the hot and dry season. 

166. If ormal Direetion 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 nonnal 
condition. (213-217.) 

§ 2. Their Structure and Forms as Organs of Assimilation 

OR Vegetation, t. 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 parcneh\*ma, and in this the work 
of assimilation is carried on. The woody is the fibrous frame- 


noi'k. the scpnrate parU or raiiiifitiitioca of which form whi 
vmiuiislj' ealleil the ribt, — a stillk-ienth' pn>per t*nn, — nerien or 
vtins. Tliu lutU;r iiuiuls muy siiggt'st false aiinlo^fs ; but tlicy 
ai* of tliu commoiK'&t uae iu ilescriptive tiotany. TUal of veiut, 
mid of it« diminutive, \-eiaUu, for the sinallvi' ramiflcatioDs, is 
not amiss ; for the fibrous framework not only gives tirmiii-ss 
mill sujipurt to tli«< softer cellular apparatua, t. e. I'omis ri])§, 
hut servi-s in tliu leaf, aa It does in tlie stem, for thv more rapid 
Con^Tyante ntid distribution of the aap. The subdivisions con- 
tinue beyond the limita of unassisUxi vision, until the ftbH>.va8- 
CuUr bundles are nnluced to attenuated fibres ramilied ttirough 
the parenchyma. In leafstalks, the wooily Ituudli^ arv purullel, 
not raimlltMl, and airanged in varioiia wajs ; iu Ksogens usually 
BO as to funn in ci'oss-section an arc or an ineompletv or com- 
plete riug. In leaves serving US foliage or oi'gaus of assimilation, 
the blade is the im|M>i1ant part, and this only is here rrgaiile.1. 

IfiH. The ehuraeteristic conteiits of these eelte of pan-iidi,viiui 
aiu gnUns of MorojihyU (US'), literally lraf-gr«tii, tu which 
the green color of foliage is wholly owing, and which may be 
rcganled as the moat iini>0Jlant of all vegetable products ; 
lieoause it is in tUeni (or in this green matter, whatever iU form, 
ISD) that all ordinary assimilation takes place. As it acts only 
under the iulluence of light, the expanded leaf-blade nmy lie 
viewed as an arrangement for exposing the lax^est practionhlo 
amonnt of tliis green matter — the essential element of vt^eta- 
tioa — to the light and air. 

lO'.i. Tin' I'lirciicliyinKMwlls. constituting the green pulp, are 
„ ^-^-^ themselves nrrangcd in accordance 

- ■" ~ with this adaptation. The up]>fr stra- 

tum is mostly of oblong cells, compactly 
",-1 arrangnl in one or more layers, their 
'■^ lunger dianieU-r peri>endleuhir to the 
( ' ,_ BUi-faee. Tlic strntmn next the lower 

"^ sorface of the leaf consists of loosely 
1 _ ~1 ^ aiTangiil cells, with lougi-r diameter 
|_1_^ - ^ usually imrallel to the phine of Ihelcnf. 

' - ' i— "" 'i often im-gular in form, and so disiJosefl 

03 tu leave iuterveuing sinuous aii-spaces iH'ely iwrmeatiiig all 


pro. i3«, A mvninea twiion ibn 

■Unaiu, nlHnrlui ilxi IrncHbi >■•■>■» o\ 
till Diiper U}H iif Ibi ttM«i |ml|>, Ihi 
pMtnl. HH tn Inra only nli>uUi viinn 
Urea unl oiplmu In t1i* mt of llie I 
>il>.' Urn vH.lmiita or tklw of Ihu diu 

ith the tlilrVnfM u[ a luf of IMIrlum Florl- 

nlK iT whlrli IpljHWl venlritllyl nn vrn n<ni- 

ti, wlisrvtho »!)■ are »i^ IrmiwJjf nmnin-l: 
rrCa] Mxl or ibo luinir *Mlii« uC tin Igof (A), 


that part of the leaf. (Fig. 136, 137.) Hence in gootl part the 
deeper green hue of the upper, aud the palor of the lower face 
of leaves. 

170. Epidermis. The whole surface of leaves, as of joiing 
stems, is iiivt'sted with a trausliieeiit membrane. com|X)sed of 
one or sometimes two or three lajers of em|»ty and ratlier- 
thick-walled cells. This is the skin or rpidermis, which is so 
readily separable from the Miiccidcnt tissue of such leaves as 
those of Stoneerop atul other species of SSedum. It is of a single 
hijer in the Illicium (Fig. 13C) and Lily (Fig. 137) ; of as 
many as three in the firm leaf of the Oleander ; is generally 

hard and thick in snch coriaceous leaves as those of Kttosporum 
and Lniinrstinns, which thcrcliy the bettor endure the dry air of 
rooms in winter. 

171. Stomala or BrealUng-pores.' The epidermis forms a 
conthnioiia ]irotcctivc invcsl)nuntof the leaf except whei-c certain 
organized oix'ninsfs occur, the siomata. They are fonned by a 
tran»fijnniition i>f sonic of the cells of the epidermis : and consist 
usually of a pair <jf cells (ciillwl guanlian-cells) , with an opening 
Ix'tween thi-ni. which comnninicates with an aii'-chaml)er within, 
and thence with the irregular intercellular spaces which i>ermeate 
the interior of the leaf. Thn)ngh the siomata, when open, free 
interclu-inge may lake place Ix-twccii the external air and that 

FIQ. IJT. A mncnlik-l »«il"ii tlimnBh fhe Ijikkiiiw at n minnlo j.h-cB of tlm 
nf tlip Wlillt [.ill nf Hit' eanlmis. BtuiKliig alsna portlin of tlia giiil«r*kle irith w 
iBvatbiTjf-por*-''*. 1^10111 nt£u 



within the loaf, antl tima tran spiral ion l>e much facilitated. 
When closed, thie intert-hangc will be interrupted or impeded. 
. The mcehaniBm of stoniata is somewhat 
recondite, aud will be illustrated in the 
anatomical and physiological volume of 
this aeries. 

172. It is only when leaves assume 
vertical or edgewise position that tlie 
stomata are in equal tiurabers on l>otl) 
faces of a leaf. Onlinarily, they occupy 
ua or most aboimd on the lower face, which 

is tiimcfl away from the sun ; but in certain coniferons trees the 
reverse of this is true. In the Water Lilies (Nymphaja, Nuphar), 
and otlicr leaves wliich 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 
iacli of surface in (lifferent leaves. In the Apple, there are said 
to be about 24,000 to the square inch (which is under the average 
number, as giveu in a tabic of 36 species by Liudle,v) ; so that 
each leaf of tliat tree would present about 100,000 of these 
oriBces. The leaf of Dragon Arum is said to have 8,000 
stoniata to a s<(uare in<'h of the upper surface, and twice that 
numl>er in the same space of the lower. That of the Coltsfoot 
has 12.000 stomata to a square inch of the lower epidennis, 
and only 1.200 in the up|K'r. That of the White Lily has 
fh>m 20.000 to C0,000 to the square inch on llie lower sur- 
&ce, and perhaps 3,000 on the upper; and they are so re- 
markably lai-gc that they may be discerned by a simple lens of 
an inch focus. 

172. Venation, the veiriing of leaves. &c., relates to the mode 
in which the wooily tissue, in the form of ribs, veins, Ac, is 
distributwl in the cellular. There are two principal mo«les, the 
paralM-vtined and the retieutiiled or nelted-vetnfd. The former 
is es|]eeia]ly etinrneterislic of plants with endogenous stem and 
monocotyledon oua embryo, aud also of gj-mnos|>ennou8 trees. 







rmli of Ibo O 


1. Daluo] 




>la«nin»l «»* or tho 10.00011. \mn « 


1.0 .'i.i.1,.1 

nil ..! 


ru«»rui<ii<iirc>f 1 

lU KuOMIk, 













vhich have exugeoous steins and at least dicotyledonous 
ombrjos. The latter prevails in ordinarj- plants with exogenous 
stem and dicutjledonous embrj'o. 

173, Par»llel- reined or Ner»«d leaves (of which Fig. 143 is 
an illustration) have a ftiiinework of simple ribs (called by the 
earlier botanists iiervtt, a name still used in descriptions), which 
mn 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 agiex. The 
Banana and Canna are familiar illustrations of a mode not un- 
common in ti-opi(.-al or subtropical cndc^iis. in which the thrcatls 
or '• nerves " run from a central rib {midrib) to the margin. 
Para ilci- veined leaves are generally entire, or at least their 
mai^ins not toothed or indented. The princii>ol exception to 
tliis occurs when the ribs or the stronger ones are few in number 
and ladiately divei^ent. as in the jtahtUiform leaves of Fun- 
palms, a iiecnliur modification of the parallel- veined type. 
Between leaves with ner\'es whollj' of basal origin, and those 
with neiTCS all springing from a midrib, there arc various grada- 
tions, and also in re9|>cct to cuniug. But parallel- veined or 
ncr\ed lca\'es may be classified into 

« Qalncf. of the nctlnl-Telned or rEUcalaleil a 

FIO. l«, A la 
bUiLo ib), petlola or 

PIU. 113. PanUel-relned leaf or U» Lll; of Iba Valley. CoonlUiU m 



Batal-ntTved, tlint is, with tlic ut^rvtss all s]>niiging rrom the 
base of tlie leaf, iinil 

Coilal-ucFTcd, sprioging Irum a midrib or coata. Either may In,- 

Efctiiiervrii, the iien'es iiiiitiitig etrsiiglit Hum origin to apex 
or margin of tlie leaf, aa the case may be ; 

Currinerred. when ourviug in their eouise, aa in the leaves 
of Funkia and in Caiiua ; 

FitUielliuentd, where straight nen'es and ribs radiate from tlic 
apex of the petiole, as in Fan-palms and the Gingku tree. 

174. In typiiyil paralleUveined loaves, all retieiihitioii is eon- 
fined to minuUi and straight i-ross-veinlcts : in many, tlieso aro 
eoarscr, brauelimg. and i-eticnlat(>d ; in some, as in Sinilax and 
Dioacorea, only the primary ribs or strongest iier\-e9 are on the 
l>arallel- veined plan ; the spaeo between being tllle<l with ivticu- 
lationa of various strengtli ; thus ]>assing by gnulations intf> 

17o. Reticolated or Ketted-velned leaves. In most of these, 
from one to several primary iiui-tions of the fjamenork arc 
pnrtieularly robust, and give origin to mueh more slender rara- 
iQuations. tliese to other still smaller ones, and so on. The 
strong primary portions ai-c Itms {fo$ia) ; the leading i-aniifl- 
(tations, Vei>s (ivdic) ; the smaller and the ultimate subdivisions. 
Veislets (venula). All or some of the veins and veinleta are 
said to anailomose, i. e. variously to connect with tlioae from 
otiicr tninks or ribs. ap|)arcntly in the manner of the veins and 
arteries of animals, forming meshes. Dnt. as theiv is no o)x>ning 
of ealibre of one into anoUier. the word is etymological ly rather 
misleading. More |iro|>erly, it is said that the veins or veinlets 
foiin reticiiiafioni or net-work. A primaiy division of reticulated 
leaves, and Indeed of nerved leaves also, into two classes, is 
founded upon the mnnber of primary ribs. 

liG. There may l«e only a single primary rib ; this traversing 
the blade from base to ti|J through ita eentre or Slx'ib (as in Fig. 
142, 152-loG) is called the Midrib. There may be others, gen- 
erally few (one, two. three, or rarely four), rising fh>m the n|>ex 
of the i>ctiole on each siile of the inidrih. running somewiiat par- 
allel with it or more or lees diverging ttom it : these are lateral 
ribs. Among parallel- veined leaves, tlic Rjinana. Canna, &c., 
\tave a single rib, tVom which tlie veins fin tlic older nomencla- 
ture here called nerves) all proceed. Moat Lilies and the like 
have several approximately parallel ribs, but the midrib pre- 
dominant: in other eases, tlie midrib is no stronger than the 
others. In Fan-pnima, the ribs are mdiately diveigent, giving 
a fan-shai)ed or rounded outline to the blade. In retieulatcil 
leaves, in which the I'cins all spring (Vom the ribs, the two 


classes into wliich tlioy divide are the pinnideJy veined and the 
palinalefff veined. 

177. Flnnatelj or Feather-Teined (or Penninerved) leaves 
- are actonlingly those of iviiicli the veins and their sntKlivisions 
arc side branches of a Hin<;k'c-entralril> (midrib), trhieh traverses 
the Wade I'l-om base to .ijwx : the veins thus l)eing dis))osed in 
the manner of ttic [iliimo on the sliaft of a feather. (Fig. 142, 
152. &c.) Sometimes these contiinie straight and imdiminished 
from mi<!ril) to margins (straight- veined, as in Beech and Chest- 
nut, Fig. 152), sending otT only small lateral veiiilots; some- 
times tliey raniit^' in their course into secondary or tertiary veins, 
and these into veinlets. l*innatc venation in reticnlate<l leaves 
naturally belongs to leaves which arc decidedly longer than wide. 

17^. Some of the primarv veins, commonly among the lower, 
may be stronger than the rest, anil thus take on the character of 
ribs, or by gradations pass into such. The leaf of the common 
annual Sunflower (Fig. loo) becomes in this way tripie-ribhed or 
tripli-nerved. The appearance of a second pair of such strength 
ened veins makes the venation quinlupU-ribbed ovijiiinlupli-nened. 

Through the apin-oximation of such strong veins to the base of 
the blade, this venation may pass into the 

179. Palmately, DlRHaleij, or ItadUtely Veined (or Paimi- 
nerved) class, of which leaves of common Majiles and the Vine 
are familiar examples. (Also Fig. ISB-ICO, &e.) In those 

Fia. iti-is;. 

explained tn tbo Ii 



tlipro are thxVQ. five. Boven, or Bomctiiiies more riba of equal 
strength, the central being the midrib, and each with its system 
of veins vrliich ramify and form mcslu-'s in tlio interspaces. Here 
the wliole woody portion of the leaf diviiles equally into a num- 
ber of i»irts u[jon leaving the |)etiule or cnt<.'ring tlie blade. The 
ribs there commonly diverge more or less in a palmate or digiiaie 
manner (t. e. like outspread fingers of tJie hand, or the claws of a 
liird, or like radii of more or k-sa of a circle) : so, in the corre- 
lation of outline and veuation, this class of vcining goes with 

roundieb circumscription. This ia not so true, however, in a 
special case, viz. , where the riba, however divergent below, ciino 
forward and all nui to the apex of the lilade, thus imitating the 
parallel- veined s^ysti'ra. as in Rhexia and generally in the family 
of which tliat genus is the single northern representative' 

180. Forms as to OalUne, Ac. DeCandnlle conceived the shape 
of leaves (botli the general circumscription and the special con- 
flgiiration) to depend on the distribution of the ribs anil veins, 
and quantity of the parcnchinna in which these were outspread, 
— a tfx) mechanical \iew, and not conformable to tlie hislorj' of 
development. This proves that the fVamework is adapted to the 
I)aronch_\-ma, which grows and shai>es the organ in its own way. 
rather tlian the parenchyma to it. It were Iwtti'r to say that the 

1 In LiiinRnn lerminolngy. jmlmolt aixl diyilale tvUmA to pwlicukr oui- 
Unc only, ami were wparmtdjr u«rd to dpluilr extent o((liti«on. — /«iJmiU(, not 
iliTideil linwn to the jieliolr, rligiiale, when dm.leil, like Ihe clawi of a I'inI, 
qujte (town tlir boac. pFCundolle i^nenilizvi] llie use of the fonniT term, 
and ever lincc the two have bven lucd interalianKobly. 

FIO. lss-t« ' 

uftonniorslnipla, cblcfl; palmBtoly reined loniM. 


two eloments of the structure are correlated. Descriptive terms 
a[>plicd to leaves are equally applicable to all expanded organs 
or iHtrts, and indee<l to all outlines. Some leading forms are 
here enumcrat«<l ; and all are defined in the Glossary-. 

181. Aa to ^neral ClrcomscripUoD, proceeding from narrower 
to broader sliajws, and then to those with either narrowed or 
notched base, leaf-blades arc 

Linear, when narrow, several times longer than wide, and of 
about the same Itreadth throughout, (t'ig. 107.) 

Lanceoiate, or Lancfshaped, when Bcvcml times longer than 
wide, and ta|>oriug upwaitla (Fig. Iu3, 168), or tapering both 
upwnnl and downward. 

OWo»j, wlicnncarlj- twice or thrice as long as broad. (Fig. 169.) 

Ellipiitai, oblong with a flowing outline, the two ends alike 
in width. (Fig. 170.> 

Oi-al, the same as broadly' elliptical, or elliptical with the 
breadth considerably more than half the length. 

Orate, when the outline is like a section of a hen's-egg length- 
wise, tlie broader end lieing downward. (Fig. 171, 155.) 

Orbicular, or Rotund, drea- 
lar in outline, or nearly so. 
(Fig. 160.) 

Obiivalt, inversely ovate, or \ 
ovate witli the narrower end 
toward the l)ase, the broader 
upward. (Fig. 175, 145.) 

Cuneate, or C'lnriform. that is. Wedge-xhapef/. broad alwve and 
tapering by straight lines to an acute base. (Fig. 176, 148.) 

Spatulate, rounded nlwve, long and narrow below, like a 
spatula. (Fig. 174, 147.) 

ObtanceoliUe. inverted lance-shaped, i. e. such a lanceolate 
leaf as that of Fig. 1 68, but with the more tapering end at base, 
as in Fig. 17.3. To those who restrict the term lanceolate to the 
sense of a narrow leaf tapering equally in both directions, the 

■ or Tsrioiu ■Impls tMr«L 


term oblanocolnte is eiipertliioiia. The following t«rma itesig^ 
nato ienvea with a notchod inelcarl of narrowod base. 

Cordaie. or J/i-nrl-t/iiiped, wlieii n k'lit' of an o\ato foim. or 
something like it, hns the outhiic of its rfnimled base turned in 
(forming u iiut<^'h or sinus) where the stalk is atlndied, as in 
Fig. 172, lot. Also Fig. 149, Pontedcria. a leaf of llic pnrallel- 
veinetl class. 

Renifurm, or Kldnfy^haptd, like the laat, only rounder anil 
broader tlian long. (Fig. 158.) 

Anriculnir, or Eared, having a pair of small and blunt pro- 
jections, or eiirs. at the base, as in Magnolia Fraaeri, Fig. 17«. 
Sagiltate, or Arrow-shaped, where such ears arc aeute and turned 
downwards, while the 
main Iwxlyof the blade 
tapers upwards to a 
Ijoint, as in the com- 
mon sjiccies of Sagil- 
larift or ArRiw-head. 
nnd in tlie Aitow- 
leaved Polygonum, 
(Fig. 1C5, ItV.) 
Ifisfa/f. or JMhtrd- 
'■' ''' '™ shaped, when such 

lolws at liie base point outwar^ls, giving the leaf the BlinjM! of tlic 
halberd of the olden time, as in Po],\'goiiinn aiifoliiun (Fig. 17'J) 
and .Sorrel, Fig. 103. 

1«2. Prilate or S/neld-ifiaped leaves are tliosc in whieh a blade 
of rounded or sometimes of other shajw is attached to tlie petiole 
by some part of the lower surface, instead of the liasal margin : 
those of ^VatiT-sliield or Braseuia, of Nelumbituu. and of Hjdro- 
cotyle umbi'llala are marked examples. 'Flic anomaly is mor- 
phologically eiqilained by a eompariaou with deeply cordate or 
rcniform leaves having n narrow sinus, snch as tliose of Nym- 
pliica or Water Lily, and by supposing a union of the nppnixi- 
mtitcd cdgi's of the sinus. Fig. 159 and IGO, from two 3|R'cies 
of Iljiflroeotyle, one wiili open and the other with closed sinus 
obliterated by the union, illustmte this. 

183. As to Extrcmlt;, whether base or apex, thei-e are several 
descriptive terms, expressive of the principal modifications; 
such as 

Jciiminale, tapering, cither gnulnally or nbniptly, into a 
narrow more or less ])rolongcil t«rmhiatiou. (Fig. 180.) 

c, anil buUIo leave 



Acute, eading iu an acute aiigle, without special tapering, as 
in Fig. ISl. 

Obtuse, eiuiing with a blunt or roundish oxtromity. Fig. 182. 

Truncate, with t<.'nnination as ircut ott' by a straight transverse 
line, as in Fig. 1»3. 

Relate, with an obtuse ostremity sliglitly depressed or re-enter- 

ing. 1 

I Fig. 1«4. 

arglnaie, with ft more decided terminal notch, Fig. 185. 

Obconlate, invei-scly tieart- shaped, i. e. like cordate, but the 
broader end and its strong notch at apex instead of base. Fig. 
18G. This and the fulluwing terms are applicable to ai>ex only. 

Miicronate, aliniptly tipiH-d witli a small and slioit point, like a 
projection of the midrib, as in Fig. lf*8. 

Cuspidate, tipt>ed with a slmr|j aud rigid point, as in Fig. 187. 


184. As to HaT^in or special Oalllne, the terminology proceeds 
upon the convenient supposition of a blade with quite entire margin, 
but anliject to incisions, which give liso to notclies or clefts, if we 
regard the sinuses ; or to teeth, lobes, segments. &c., if we regard 
the salient portions betn'cen the sinuses. Ttic riiis. or the stronger 
veins, &c,, commonly tenninate in tlietwth or lobes: butinCieuta 
mncutata. and in a few other cases, they nin to the notches. 

185. Dentation relates to mero mai^nal incision, not extend- 
ing deeply into the bhtdc. The blade is said to be 

Entire.' when yj. />,/); 

themai^niscom- AA,-\f\ '"' 

pletely fillt^l out r "^^J -V .n 
to an even line, as 
in Fig. 173-179. j' 

Serrofe, when J V; 
with sm.ill and I 
siiai-]) teeth direct- 1 
ed forward, like \ 
the teeth of a saw, \H, 
as in Fig. iny. w A 

Semilate is the in 
diminutive of sen-.ite. and is equivalent to miuiilely serrate. 

' liileijirriinm-(i-'im, or ipiil-- luliir, it the ttriii in Lntin teniiinulugy. lilt 


Dmlate, or TooChed, a general term for tootbing. speciallj- 
npplietl to the case of salient teeth whitrh are uot directed for- 
ward or towards the apex of the Iiladc, Fig. 190. 

iVcnale, or Scalloped, the same as dentate or serrat«, but with 
teotli much rounded, Fig. 191. 

liepaiid, or Undiilutfy when the margin is a wavy line, bending 
slightly inward and outward. Fig. I!t2. 

Hinnale, when this wavy line is stronger or distinctly sinuous, 
as in Fig. VJ'i. 

Incised, when cut by shaip and irregular incisions more or less 
deeply, Fig, IDJ. This is intermedials between dentation and 

mti. Lobatlon or Segmentation. When the blade is more 
deeply penetrated by incisions from tlie mai^n, that is. when 
the spaces between the riba or principal veins are not fiUeil to 
near the general outline, it is said to Ijc liAed, elrji, parted, or 
divided, according to the degree of separation ; and tlie portions 
are ealle<l loliei, tegmmlt, diciiiont, &q. The most general name 
for sutli parts of any simple blade ia that of lubv$. More par- 
ticularly a leaf-blade, or other body, is said to be 

Lobed, when the division <^xl«nds not more than half way down, 
and either the sinuses or the lobes are rounded ; 

iieft, when the division ia half way down or more, and the 
lobes or sinuses narrow or acute ; 

ParUd, when the divisions reach almost, but not quite, to the 
base or the midrib : 

Divided, when they sever the blade into distinct parts, which 
makes the leaf c(nn^oun(/. (Iil3,) 

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 di\ided or parte<l and 
these primary lobes again lobed or eleft, the lobes of 6r8t onler 
are commonly called ISenuEVrs (sometimes dirition$ or partiliom), 
and the parts of these, Lobft, Or the lobes may be designated 
as primarj'. secondary, tertiary, &c. Ultimate (Ktrtioiis or small 
lolwa may be called LubuUt or JjHieltU. Also tlie iwrtions of a 
quite divided blailc take tJie name of Leajktt. Hy jirojier selec- 
tion of terms, the degree of division or lobing may thus be 
esiiiresaed in a single wonl. 

1S8. As to Number of ports, tliis may bo teraely expressed by 
combination with the adjective term applicable to the degree ; as, 
Two-tohed, Three-lobed, Ftee-libed. Many-lobed. &c. ; or Twm-Five- 
clrf\, Mann-clefi. &c.. in Latin form Blfd. Trifid. Afubifd, &c. : 
Ttro-Jivf -parted, Ac., sccortliiig to the nnmlter of dirialons 
which extend almost to the base or axis ; Tvio-Fivt-divided (in 



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 ma}' be simpl}- and best 
expressed by taking into account the nature of the venation or 
the distribution of the ribs, &c., which controls or is co^oixlinated 
with the disposition of the lobes. Pinnatel}' veinecl leaves, 
when lobed, must needs have the incisions directed to the mid- 
rib ; palmatel^'-veined or radiated, to the ai)ex of the petiole. 
The lobes or dinsions of the first will be pinnatel}', of the second 





palmately disix)sed. Acconlingly, the three leaves of as many 
species of Oak, Fig. 195, 19G, and 197, represent respectively a 
pinnafely lobed, pinnately cleft, and pinnately parted leaf, while the 
accompanying leaf of Celandine, Fig. 198, \% pinnately divided. 
The first three, however, when the degree of incision is not par- 
ticularly in question, usually pass under tlie common term of 
pinnatijid. Fig. 195 nuKlcratcly, Fig. 197 dei»ply. The number 
of lobes, when definitely marked, may come into the descriptive 
phrase, as pinnately 1-lohed, pinnately 1 -cleft, parted, or divided, 
as the case mav be. 

190. Similarly, Figui-es 199 to 202 represent, respectively, a 
pahncUely three-lohed. three-cleft, three-parted, and three-divided, or, 
in Latin form, trilobate^ trifid, tripartite, and trisect or trisected 
leaf. Fig. 166 is vl palmately b-parted leaf; Fig. IGA, palmcUely 

FIG. 195-198. Pinnately lobcd, cleft, part€<l, and ilIvUlwl leave*. 
FIO. 19e-2U2. Palmately 3-lobed, cleft, parted, and divided leaves. 



mutlifid, &c. Fig. lt'>2, a leaf of Dr:^on Amm, ia palnn 
9-paried. But, as the InternI sinuses are not so docp as the 
otljere, the leaf is said to be pfdate/i/ parted, or pedale, iu tlie 
early terminology. 

191. Moreover, as the lobes or divisions or a leaf may be 
again similarly lobed or parted, &c., tiiis composition may be 
indicated hy the prefix twice, tlirice, Ac. as ttcice pinnalljid or 
hipianalijid, tkriee jiinnaUli/ parted, thrice palmalfit/ parted, and 
tiio like. Thus, a wonl or two. or a short phmse, Inay tiescribc 
even a complex leaf, so as to convey a peifectly clejr and defi- 
nitL' idea of its conformation. 

182. 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 
ltae«tf which have l>een tjius far considered, have a single lamina 
or blade, which may, however, at one extreme be entire, at the 
other inany-]mrted, and even several times tlivided. 

193. ('ompotmil LeiTes are those which have from two to 
many distinct blades, on a common leafstalk. Tliese blades, 
calli-d Leaflets, may be sessile on the comuwu kufstitlk, or they 
may have leafstalks of their own. As the leaf very commonly 
BC|>aratcfl in age by an articulation of its petiole with the stem, 
so leaflets are commonly more or less articulated with Ihc com- 
mon petiole. When the leaf, witli its {>etiole, falls fVom tlic 
stem, the leaflets may as completely separate rw>m the L-ommon 
petiole. They do not always do tlits. Divided leaves, such as 
Uiosc of Fig. 138 and 302, though ranked among the simple 
sorts, are com|>ound in the sense of hating distinct blades, 
but without articulation. Some of these blades are a[it to be 
confluent : tlial is, a divided leaf is often in [jarl merely partcti, 
as ill the upper portion of Fig. 198. Such leaves ai-e so intcr- 
mnlitito between simple and comjiound that it btKHimes indiflbr- 
ent. or a matter of convenience tu be settled by analogy, under 
which head or by what language they shall Ik? dcscrilwd. How- 
crer. most leaves are so constituted as to leave no doubt whether 
they ai-e sunple or comiwuud. 

194. The leaflets of a compound leaf being homologous with 
tiie lobes orscgmcntsofa simple leaf, indeed being such segments 
hilly isolated, the two sorts fall under the same tyjjes. A [lin- 
niUi-ly velnt^l simple leaf is tlie homologue of one kind of com- 
[xiund leaf; a railiately veiued leaf, of the other. TItat is, 
compound leaves are either pinnnt* or palmate, 

1^5. Pinnsle Leaves (Fig. 203-205) are those in which Uio 
leaHels arc arrangetl along the sides of a petiole, or rather of its 


prolongation, the RuACms, which answers to the midrib of a pin- 
natcly vcineil simple leaf. There are three principal sorts, and 
some snbonlinate ones. That is, a pinnate leaf may be 

Impari-pinnate, or pinnate with an odd leaflet, t. e. a terminal 
one, as in Fig. 2U3 ; aiul this is the connuoner ease. 

Oirrfiiferout P'mnale, or pinnate with a tendiil (Fig. 204), as 
in the proper Pea triljc anil Bignonia. Here either the termi- 
nal leaflet only, or the upi^er lateral leaflets uUo, ore ifplaeed 
by tendrils. 

Piiri'pinniUe, or Abruptly Pinnate, destitnte of a terminal leaflet 
or of any thing answering to it, as in Fig, 205, 

Interntpledly Puinate denotes merely a striking incqnality of 
size among the leaflets : Lt/raiely Pinnate, one id which the termi- 
nal leaflet is largest and the lower small. 

196. Palmate or Digitate Learcs (Fig, 206, 93) are those in 
which the leaflets all stand un the 
summit of the iK'tiole, Drgliate 
(lingered)was theold name, when 
the term ptdmnte was reatrictitl to 
a simple but iwdniatt-ly lobeil leaf 
of this tyi>e. But since the time 
of DeCandolle the two names have 
l>een useil intCR'huiigeably, Pal- 
mate leaves have no primary dis- 
tinction into sorts. exwi>t as to 
the numtier of leaflets. These can 
never be very numerous ; but there are fully a dozen in some 

FIG. va. An Impnrl-i.iniiatu or ixlcl ijinnUe leaf. SM Plntiato viLh % Ittnilrll. 
90S. Atim|AlyflnnBtuleBfuraCa»J>. 

FlU. 200. Pklumlelx or JlgllaKlf S-fuLIuUt« le*f of ■ Buckeye, JEm't! 



; only five to nine, or'ral^l 

More commonly there s 
. rarely two, or econ a single ( 
l'J7. Number of leaflets may In? indicated by an adjoctice 
expressitin comi»sed of iUp proper Latin numeral prefixed to 
fotiolale {Foliotum, diminutive ot folium, answering to leaflet). 
Tliue, bifoliolale, ot two leaflets; trifoliolale, of three leaflets; 
guadri/olicilate, of four ; quin^urfoliolate, of five : plurifoltolate, or 
mi/Jti/ii/iolalf, of several or numerous leaflets, &c. Tbese tcrma 
are still more deseriplive when accompanied by the word pin- 
nattly or palmat^ly, inilicative of the kind of compound leaf; as, 
palatatflg or digitaUly trifotiolate (common Clover-leaf, Fig. 211), 
or S^u/io^fe,as in Uuckeye(Fig.206), andaoon. Also, pinjiateig ' 
i6-/olwlale, as in Fig. 205, or 17-foUolate, 08 in Fig. 203; 
pinnattly trifo/iolatt, as in Phaseotns, and in the low Ilop-Clover, 
Trifolium procumtwna.' 

198. But, in either class of compound leaves, the leaflets may 
be reduced to a minimum numtier. A pinnately trifotiolate leaf 
is one of the impari-pinnatc- kind reduced to three leaflets, to one 
pair and the odd one ; and thia is distinguished tVom a palmatcly 
trifoliolatc leaf by the attachment of the pair at some distance 
below thi! apex of the ix-tiole, and by the aiticulalion above this, 
vhieh marks the junction uf the terminal leaflet's petiole (or its 
base, if sessile) with the rhaehis or common i>etiolc. 

lUU. UnifvUoIate compouml leaves (by no means a direct con- 
tradiction in t«nn3) are by this articulation distin- 
guishc<l IVom simpL' leaves which they simulate, 
' See the leaf of the common Barbciry, Fig, 207. 
t In other sitecies, of the Mahonia section, the leaves 
; are all pinnat«ly S-O-foIiolate. witli well-developed 
' common petiole : in the true Berlwris, they are all 
thus reduced to the terminal and long- petiolu late 
leaflet, on an almost obsolete petiole. Orange and 
Lemon leaves are in siiuilarcase, lint with the joint 
close to the blade. A «im|jarison with near ivla- 
' lives shows that tJiesc are also iinifoliolate leaves 
of the pinnate kind ; though tliis could not be ascertained by 

200. Deconponnd or Twice and Thrice Componnd Leaves. These 
are to once pimiatc or once pnlroate lea\cs what the hitter are to 

> Id piiinntc leftvci, encli IcnHct usunlly lias its appu«ll0 fi>11ow. and (he 
numtier may be indJoI«l by tlie pain, ni amjiyait, UJiyiiif, Iryjagnlc, ntiil 
pluiiiagale, accunlioK to tbe number oljityii. m pnin. 

Flo. XT. UliinillnUtelaiirnf Berbarif vulK*rl>, wIlhlKuUuI iwUule utluulaleil to 



simple leaves. Aa 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 onler. Decompound is a 
good geueral uame for all more than once comiwunded leavea ; 

but the name has been applied rather to iiregxilarlj many-times 
parted or disSLCtcd tea^es (such as those of DicoDtru) or to 
those more thin thriLC compouude<l Of reguhirlj twice or 
thrice compound leaves the commonest are the 

BipinnaU or Tu»i.t Pinnate ol orliiniv 
occurrence m the Miinoscous and tiiaaliii 
neous, but not m thr 1 apilionaceous Lcgii 
minosie Fig 208 rtprcseiitsabipinnitc Itaf 
of the Honej Locust ((..leditschia), with the 
variation (common witli that tree) that some 
of the i>artial {>etioles. in this figure only the 
lowest, hears a single leuflct, while the others 
arc cxlende<I into secondary rhachisea fur- 
nished with numer- 
ous leaflets, mattly 
ID the abruptly pin- 
nate style. On the 
same tree, the earlier 
leaves, wlijch j 
clustered on short , 
spurs, are siin|ily ' 
pinnate. The largo 
loaves of GjTiinocladus are similarly and abniptly bipinnate, 

FIG. JM, Blliin 

a leave* of Scu'U 

>r niaiLtiwiiiii 111 

at, with >i 



1 is simply {jiunatu or nilh one or two 

except at tlie haec, wliii 
pail's of simple leaflets. 

TripinnaU or Thrice Pinnate leaves of a regular sort are rare ; 
but. with sfniie irregularity, lliey occur iu many sijecies, as in 
Aralia, &v.. Tliis extent o£ di\'isiou, and even mucti greater, is 
common in Fenis. 

Digitate-Pinnate is where tlie primarj- dinsion of the petiole ia 
on Iliu iialmate or digitate plan ; the sccoodnry, on the pinnnt«. 
Tbis seems to be the ease in the Sensitive Plant, Mimosa pudica. 
Fig. 209. But tJie leaf is here truly bipinnato with the primary 
divisions very eroM-ded at the apex of petiole. 

ConfugaU-Pinnale ia the same arrangemeut, vith the primaiy 
divisions a suigle pair, at the a|K'x of tlie petiole, and the leaflets 
pinnatvly arranged on these. 

Digitatelg or Paimalelg Decompound in a nearly regular way 
j^ is not an uucoiuniou cose. 

1^^- Usually, the (jetiole is siiL-ees- 

sively tliree-forkeil, as in Fig, 
210, wlien the leaf is said to 
Iw biUmate (twiee tcmatc), 
Irilernale (llirice temate), or 
quniiriteniale (four times ter- 
nate). &v.. aLiiocding to the 
number of times it divi<Ie8, or 
The ultimate divisions in audi 
caees of threes arc commonly of the pinnately trifoliolnte tj'pe. 

201. Finnic is a eonrenient name fiir the partial ]>etiole8 of a 
bijiinuatc leaf, taken tt^etlier witli the leaflets that l>elong to 
tliein. Thus, the 8ensitive Plant, Fig. 209. has four pinuie, or 
two pairs ; the Honey Loeust. Fig. 208, a greater number, 
Wlien BiieU leaves are still further comgioundeil. tlic pinnae of 
higher onler, or the ultimate ones, take the diminutive term of 
PiXKiLX or Pis^LLEs, The blades these bear are the Leaflets. 

202. The Petiole or LeafNlalh is a eomparatively unessential 
part of the leaf. It is often wanting (then the blade is seuile) ; 
it may Itc absent even in compound leaves of the {lalmatc tyiw, 
the leaflets rising side by side from the stem. Wlien present, it 
is UBimllj' either round, or half-cylindrical and ehannelled on the 
npper side. In the Aspen, it is flattened at right angles with 
the bla<le. so that the slightest breath of air puti the leaves in 
nmtioii. Sometimes it is nmch dilated ami menibranneeous at 

Fia. ■no. Qiukdtt-Unulclr conpauiiilM Icnialcb'decuiuiHmiiil lu&r uf TtutUcimni 




base, aa in many nmbelliferotis plants ; eomotimes it forms a 
sheath, occasionally it is boixlcred with apiHtnilages, &c. Peti- 
oles may assume siwcial functions, to bo liereafWr considered. 
The woody and vascular tissue runs lengtkwise througb the 
petiole, in the form usually of a dcftiiitc number of pai-allcl 
threads, to be ramifie<l in the blade. The ends of these threads 
are apparent on the base of the leafstalk when it falls off, and on 
tlie scar leJl on the stem, as so nianj- round dots (Fig. 81, 85, 
91), of a uniform number and arraugemoiit in each species. 

a03. PortlBl Petioles are the divisions of the iwtiole in a 
com|)ounil leaf. The footstalk of a IcaSct taiies the diminutive 
name of I'etiolule. 

204. Stipnlcs (157) are lateral apiicndagcs, one each side of 
the base of the petiole, sometimes Ciee 
lix>m 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 op|>OHitG to it, but their normal ^^ i 
position is 8up|)oscd to bo latoi-al or ^ 
marginal to the petiole. Sometimes 
they are foliaceous in apiwarauco and - 
in flinction ; sometimes they are dry 
and colorless or scale-like, reduced to 
mere e|iidermal tissue, and evidently 
fimctionleBS ; sometimes (as in filag- 
nolia, P"ig. 81, Kig-ti-oe, and Iteeeh), 
tliey ser^■e as hud-scales, and fall when 
the leaves develop ; sometimes they arc 
reduced to a mere bristle, or take the 
form of a spine, as in the Loenst (Robini 
expansions or wing-like mai^ins of the base of the petiole, 
siieh as those of tlie Sa-tiTrage tiilKi, and stipules adnatc to the 
mar^ns of the |)etiole. as in most Rosacea?, there is no clear 
limitation. But |)re3enee or absence ol" stipules gencrallj' mns 

FIO. 211. C1nTeT-lcar,wllUiulnii>c«l]ii>l«. 212. 0.'lircnt«ii)i|>ul«i(<>cliniaor«-rca| 
nf Puljrgnnaiii orlentalc, ibeUliIng llio lUiiii for BomailliUiicu.audeiidlDgi^ "'• 

lug border. 

Between salient 



Uiroiigh & natural order. Yet what are eallw! stipules in one order 
may [lass for oxpanBioim or a|)|M>iKlages of the i>otJole in another. 
Til S|K'rgiilaria, eome stipiik's nre eonnnte nruuml tlic b.ise of tlie 
pair of leaves, iiidLKling tlietu as well as tbe stem iu tiii; sheath.' 
iOo. Stipules, whieli are nonnaUy a pair, may unite into one 
liotly, either adnate to the Inner faee of the leaf, as iu some 
speeiea of Potanu^eton, or iinilcil opposite Ilia leaf, as in Plane- 
tree, or uniteil ittlerte in a slioalh, as iu I'olygonum. AIsw when 
Ihe leaves are opposite and the stipules thus hrought into prox- 
imity, the ndjaoont half stipules of tlie two leaves may eoalesoe, 
iiud present the u]tp<'arancc of only two stipules to two leaves, 
lis in many Ruhiactw. A notch or fork at the ai>es oflcu 
iiiitieatcs the composition. 

206. Sheatliing stipules, like those of Polygonum (Fig. 212). 
are said to be ochrenle, or (better) ocreale ; the sheath, thus 
likened to a leggiu or the leg of a boot, is au Ot'UKEX, as written 
liy Linnrcus. or better Ocbba. 

207. The LicuLE of Grasses (Fig. 150) is seemingly a thin and 
scnrious extension of the lining ta the sheatli which answers to 
petiole in such leaves: it projects at tlie junction of the sheath 
and hlade, there fonnhig a kind of ocrea ; and it is generally 
ri'gnnled as u sort of stipule. 

20!^. Stipels {SUpe/ta) arc as it were stipules of leaflets, which 
are common in certain tribes of Papilionaceous Leguminosa\ t. y. 
in the Plioseulcw, In Wistaria, Locust. &c. ; also in Staphylea. 
Tbey are small nnil slender, and, unlike stipules, they are single 
to each leaflet, except to the terminal one, wliieli has a pair. As 
leaves fiirnisliLiI with stipules are said to be ttiputnie, so leaflets 
with stiiK'ls are xtipellaie. 

200. Home DUDSual mmlificalions of learn as foliage. In 
leaves as illustrnteit thus far, it is the lamina or blade which is 
esponiled to do the wock of fohagc ; which is expanded hori- 
zontally, so as tj> present npiwr and under surfaces, one to the 
sky. tlie other to the ground ; which is bilaterally sjmmctrical 
or anhatantlally so, the two lateral halves being nearly IT not 
quit« alike: anc| which is afHxedto tbe stem at the basal margin, 
or some part of it, with or without a petiole. Various de^-iations 
or api>arent deviations from Uiis pattern occur. Some of them ara 
of eoiniiuratively small account aud simple explanatiou, such as 

210. InminllKterol Leaves, being unsymmetiieal by the much 
greater development of one side. This is illustrated in the 
whole genus Begonia (as in Fig. ICl), consisting of many s^k.'- 

1 Ai pointvd out \ij Ynt. A. Dickion, in Nature, xrlii. MT. 



cies, some of which are moderately, and most of them Blrikiagly, 
oblique in tliis way. Elm-leaves, and the like, are more or lees 
in^uilateral at the base. 

211. Connat« and Perfoliate Leares. These are explained by 
the union of con 
ttguous leaf edges 
Peltate lea\ es to 
whitli a iiaragrnph 
has alreadj been 
gi^cn (182) come 
under the same 
head the secmmg 
attachment of the 
petiole to the lower 
fjce of the blade 
btmg the result of 
a congenital union 
of the edges of the 
sinus In a sessile 
leaf when such a 
union takes place, it surrounds and euclosca that iwrtion of the 
stem; which \at\i\ia perfotiate. (Fig. 
213, 214.) It is the stem which is 
literally perfoliate, i.e. which seem- 
ingly losses through the leaf; but it 
is ciiatomarj, though ctymologically 
absunl, to call this a iwrfoliatc leaf I 
Uvularia jwrfoliata (Fig. 213), in the 
later groMh 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 conlate and 
clasping ; the uppermost simply ses- 
sile by a rouniled base, Baptisia 
pcrfoliata (Fig. 214) is a more 
strongly marketl ease of jwrfoliation. 
But there ore good morphol.jgieal 
reasons for inferring that this seemingly simple leaf consists of 
a pair of stipules and a leaflet combined. An occasional mon- 
strosity verifies this supposition. 

FIO. Jia Ijnts branch nf C 
FIO. IM. Lcsr^snilllnwerii 
FIO. US. Lonlcem llavk, k 

I or BaMlt 


212. When leaves ore opposite, the pcrTolintion (sncha: 
of Uoncj'suckka, fig. 215) is obrioualy the rpsiilt of a congcai- 
IaI union of thv bases of the pair by ttieir eoutt^ioiia uiges. 
Lcavca ainnale in thin way \>y oairon' 1)as<^s art; not rare nor 
nniarkuble ; but when the two arc thus coaltisceot into one broad 
(bltflveous body, giving tliis appearance of pcifoliatJon, the t«rm 
coHnalt-per/uliate is used to exi>iVBS it. 

213. V«rtle«l Learn, those with blades of the ordinarj- ktud, 
hut presenting tiieir edges instead of their faces 
to ttiu earth and sky. or when erect with one 
edge diruoted to tlie stem and the other away 
&Oin it, are not uncommon. Tbey prevail in 
the Anstralian MyrtBceic, &c.. and occur with 
leas constancy ia the Califuniian (lanzanitas, 
snd in a great variety of herbs and shrubs. The 
anomaly involves no exception to the rule that a 
leaf-hlade is always expanded in the horizontal 
plane, when expanded at all ; for, except in cqui- 
taot leaves, it is the rc«mlt of a twiiit of the petiole 
or of the blade itself.' In strongly marked 
cases, or In most of them, the organizalioii of 
the epidermis and superlteial purenchyina and 
the <)istHbution of tlie utoiiiata are tbc same on 
both faces. 

'ill, EquitaDt IrfBVOs are vertical on a diflbrent 
plan. 'I'hey are (.■onduiillcate, t. e. are folded ^ 

together lengtliwise on their middle, the upper surface thna con- 
cealed within, the outer alone [iresented to the air and light. 

' Silpliiiim lucinijttiiln. till! mwnllod Cornpus* PIbuI, and (hnrdl)- leMBof 
8. liTi-liliilliitiuufutii. arc kwhI instnm-oa of the kind, must >it llic Icnvcs 
makliiK a haif-iwiit. the nullcnl unvt by thoir long iH'tioli->. In tliu foriiwr 
vpccic), tlic pinnat«ly parted blade oL-ciHionally makes a faKlicr twiit, bu u 
to bring tlw ujipcr part inlo ■ plane nt right angle* to tliv lower. I'lic 
hiadra place tl>en»«lviia in Tarioaa dlrvctioni aa retpecla tlie cnrdjnal point* ; 
bnt i)n ilw pr«lrlM Uio grvati-r unmliBr affi'ct a nonli and nouih direi-tion of 
their edgos. — a pi-cullarily Ibat pointed out, in the year IHU, by Ueneral 
B. Alvord, U, S. A. 

HoT It1>. 

!• TontJilnrlt. 



Being two-ranked ami closely crowdeit, the outer ones at their 
base fold over or bestride the inner (na shown in tiie seetional 
diagram, Fig. 217), wlitince the name of eqitilaut. Above, tlie 
contigiioiia halves of the inner face oongonitally cohere, and so 
pntilnee the swonl-shaped or linear vertical blade which is 
characteristic of Iris (Fig- 210) and tlic Iris family. In moat 
there is a farther complication, of an cxcep- 
liLiiial kind, viz. the development hacknnnis 
of a ixjrlion of blade iVom the midrib, often 
forming most of the upper part of such leaves, 
whii'li tlierefint! may really be said to develop 
in the vertical plaiie, 

21o. Leavett with no distinction of Paris, /. f. 
of blade and petiole. Tliis is tlic case in Iria 
(Fig. -21C), Daffodil, tlic Onion, and perhaps 
of most parallel- veined leaves of Endogcns. 
Tliosc expanded in the horizontal jilane may 
however lie regarded oa sessile blades: those 
which are not exj^anded, but lilifomi, or nee<Ue- 
shajM'd (aeicular), or awl-8ha|)ed (tabulaU). may 
be regardiil either as homol(^ous with iietioles, 
or as nnexi)nn<Icd blades, which nmounta n<'arly 
to tile same tiling wiierc there is no ti'aee of a [H-tiole at base. 
Under this bend may be ranked the leaves of I'iiics (Fig. 24«) ; 
also both the subulate and the 
s<ale-sliaiwd and ailnate leaves 
of Ariior VitjB, Red Cedar (Juui- 
l>emM Virginiann).and otlier ta-es^ 
of the Cypress trilie. (Fig. 218.) 

210. Stipnips §en'in; for Blade. 
Lathyrus Aphaca is a gwHl in- 
sbmco of this (Fig. 2111) ; tlie 
IK'tiole becoming a tendrii, the 
leaflets which its i-clalivcs 1>c:ir 
tx.'ing wholly wanting, the ample 
fuliaceoiis stipules assume tlie 
api>earancc of leaves. In some 
other s|Hfics of Latliynis, and in the Pen, equally large stipules 
share with the pair or paii-s of leaflets in the ^mctioiis <)f foliage. 
On moqibological evidenc-e, we judge that the singular leaves of 

riG. !I9. Ijith' 
at ■ pair of fuUac 

at Arbor Vllip. irltli bnlli ikwI-Klinj-cl bihI mnle-Kliapi'il Ivar 
la Ai.ham: porlion if item. Warini sKiuRle Iraf. wlildi vnii 
li Mpalea, kuil k pellola In llw tuna of > undrll; In lu a: 


Baptisia pcrfoUala, shown in Fig. 214. are not simple blS 
bat eacb a pair of stipules, with or williout a tonuiiial k'aOct, ail 
completolv confluent into one Ixxl.v. Tlic relaU-cI spctit-s of the 
genus have trifoliotate leaves and Toliaivous stijiulea ; hence lliese 
simple leaves without stipules are best cxplaine<l in Iliis way. 

217. Fhjllodla, or Petioles serTing for Blade. Sometimes the 
petiole develops foliaccoiis margins, or nings, as in tlie Bitter 
Orange and in Rhns coiiallina. These are elHclent as foliage in 
proportion to their size. These are not to Iw confoiindi*(l with the 
ease in wbieli a i<etiole specially develojjs as a Made-like organ, 
which usuqis the otilec of foliage. A ))etiole-blado of this kind 
is iiariietl a Pim-Louiuit. Occ-urring only in Exogens. pbyllodia 
ore generally distinguished trom true bUdes by the parallel 
venation, and always by their normally vertical dilatation ; i. a. 
they, 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 SIXl Bjiccies. The 
true lamina of these ia bipiunato. It appears on seedlings, and 
occasionally on later growths. Several South Anieiicans|K>cie9 of 
Osalis produce phyllodia. So likewise do our tubular or tnimijct' 
Icavwl species of Sun-acenia in Uial i>ortioD of the foliage whii-h 
develops the pitclier imperfectly, or not at all. Indeed, all 
Sarraeenia- leaves are tiliytlodia wilh the back in most of them 
hollowed out into a lul>e or pileher; and the terminal hood 
i to the blade. 

5 3. Leaves sEnvtso Special Offices. 

218. Leaves may serve at the same time botli tlieir ordinary 
and some sjicciul use, or even more than one B[»ecial use. For 
example, in Nepenthes (Fig, 222) there is a well-develoi>ed 
blade, usually sessile, whieli serves for foliage, a prolongotion of 
its tip into a tendril, which ^e^^■cs for climbing, tlien an extraor- 
dlnai^' dilatation and hollowing of the oi>ex uf this into a pitcher 
for a verj- a|H-einl use. and a peculiar di'vclopitient of the aj^x 
of tliis into a lid, closing llie orifice dining growth. Among the 
siwcial purposes which leaves snliserve, and tlie studj- of whidi 
connects singulurilies of morjihology with teleology-, tlie most 
remarkable is tiiat of 

lM9. Lcufra specialised for tlie UUIIsatloD or Animal Matter. 
This occurs in leaves which also assimilate, or do the owlinary 
woi'k of vegetation ; and the sj;)eciul ftinction is usually taken up 
by some particular ijortion of the organ. The detiiila of tliia 


subject — which has of late become highly interesting — belong 
to pbysiolc^y, and therefore to the following volume, to which 

all historical references are relegated. Only the mon)bolt^- of 
such leaves is here under consideration. 

220. As Ascidla or Pitchers, vessels for maceration, &c. These 
occur in several widely different 
families of plants. The commoDcst 
are those of the Sarraccnias, natives 
of Atlautic North America. Thoy 
arc evidentlj- phyllotlia (217), the 
canity being a hollowed dorsal \kit- 
tion : the wing-like or foiiaceous poT- 
tion. always cons|iicuou8 and fonning 
the ventral border, makes the nholo 
organ or most of it in the earlier 
leaves of the tubular sijccies. The 
pitchers of S. pHr[)urea {Fig. 221, 
225) , the only sjiceies which extends 
north of Mrgiiiia, are o|)en cups, 
half tilled with water, mneh of wliieh 
may be rain, in which abundance of 
insects arc usually undergoing macer- 
ation. In S, variolaris (Fig. 22G), the hootled summit, answer- 
ing to the blade of the leaf, arches over the mouth in such wise 

FIO. MO. PHchBranfHeHflinr'hora; 221. orS«rnui!rilai.urpun;»i M2. nfXepentliw. 
S3. A pliyllodlani u( ■ New Holland AcacU. 21'4. Tlie uiuv. buailng ■ reducml cum- 
poniHl blade. 

FIO. 22a. PitcIwT-lmTe* orSuTWenla |>nrpam; ow uf tkemwltli the upper part 




lis to mostly escUide tho rain; in S. psittacina (Fig. 227), the 

iiill<.'X<.'i;l and inRatcd hood completely excludes it. The watvr | 

winch these contain is undoubtedly a si-cretio 

All entrap flics, ants, and various insects, wliich . 

in most species arc lured into tlic pitcher by a I 

sweetish eccretiou around or at some part of the 

orifice.^ Few that have enlerwl ever eseatie ; . 

most are decomposed nt Uie b<ittora of the OB\Hty. 

Ill Darlmgtonia Californica (Fig. 238), the Cali- 

fornian teprcsentative of Sarraccnia, tho inHntcd 1 

IkkhI giianU against all access of rain, while the 

orilice is freely 0|)en to flying insects from t>c- I 

ncath : and a singular two-forked a|)i)cndagc, like 1 

to a fish-tail (probably the homolt^ue of tlie J 

blade), overhangs the front. The inner face of [ 

tills appendage is besmeared with the sweet and I 

viscid secretion which allures insects to the oi>en- ] 

ing. In this and in Sarracenia variolaris, the | 

sweet secTction in the early season is continued 

ujion the erlge of llie wing, forming a saccharine 

trail whiuh leads fVoin near the ground up to the j 

orillco of the pitcher.* Fig, 220 represents pitchers 

of Ileliampliora, a, little-known South American 

representative of Sarracenia. Its wing is narrow 

and inconspicuous, tlie mouth widely o|jcn and 

directed upward, and the hood rctluceil to a 

minute and upright, probably functionless «(>- 

aa pcnduge. In Cephalotus — an anomalous plant 

of Anslralia, of uncertain afll- 

nily — the leaves for foliage 

arc dilat^vi phyllodia; among 

them are others completely 

transformed into stalked and 

short pitchers, with tliickciicd 

rim and a well-fitting lid. 

hingetl liyone edge. Fig. 229. 

The particular morphology of the parts is not well made out. 

' This BWMt secretion, which »t limps is Tory obvious in the soulhcm 
species, hu nlao l*i-n delecled bj Mr, EJwaril Itilrgcii! in S. purpurea ; liul 
il i» mrcly scon, bdiI probably plajf* no imporuiit p«ri in ihe capture and 
drowning i>f ihe mullltuile of Insects which the»e pitrhers nre spl to cont«n. 

■ Tliis trail was diwwvcred by l)r. J. H. MriUiihamp, of South Curolliui. 
See Proc. Ani, Association for Ailrancrincnt of Sckncp, nxlii. 113 (1871). 
Fia.lae. rilcliucurSarisronLiiTululmtls, £!T. Sameors. i»lLtacln*. 


221. The pitcher-bearing leaf in Ncpenlhes haa been referred 
to (218, Fig. 222) : of this tliere are various st>ecie8, all of them 
some V hat woody cli mb i ng 
plants of tronical Asiatic 
and African islands of ; 
the southern hemisphere, 
some of them familiar in 
C0118C n'ator y cu Iti vation. 
Here tlie tendril may be 
regarded aa a prolonged 
extension of the midrib 
of the blade,, and the 
pitcher, with Us hinged 

lid, as a peculiar development from its apex. Tlic 
wat*r contained in the pitcher is a secretion, ; 
of which appears before the lid opens ; and a swcclinh 
excretion at the orifice hires insects. The presence 
of these in the pitcher increases the watery secre- 
tion in which the animals are drowned ; and this 
sec-retion Is ascertained to have a certain digestive 

222. The aquatic sacs of Utricularia or Bladder- 
wort are diminutive ascidia, always under water, 
and with lid ojiening inwanl, like a valve, preventing the exit 
of minute animals cnlrap|)e<1 therein.' Moii>hologically, thcj' 
are doubtless leaves or (wirts of leaves. 

223. As SensillTe Flf'lraps. The leaves of all species of 
Drosera or Sundew arc I)eset with stont bristles tippe<l with a 
gland, which secretes and when in good condition is covered 
by a drop of a transparent and verj- glairy liquid, sufHciently 
tenacious to hold fast a fly or other small insect. Adjacent 
bristles, even if not toxidied, in a short time Wnd 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 ixitiolc and blade. In D. rotundifolia 
and other common round-leaved species, there is a clear disttno- 

' Thii wai flr»t mncte out hy J. D. Hookf^r. and announcoil in his iiWreBS, 
■« Prwiilmt of Ihe BrilUh Aeaocialion for the AdTanceincnt of Sticncc, at 
Edinfaurgli, 1RT1. 

' Darwin. IngecliTurom PTanti, Wii. Oohn, BeitrSRc tar Biologic der 
Pflanzen. 1875. Mra, Tnat, in The Tribune, Sew York, September, 187i, 
and Gard. Chron. 1875, SKi. 

FIO. 2S. FtUIicrotDaiUDEloniaCallfonitca. 229. PItcbar of Caphaloliu Ibnica- 
1*H>, witb ltd [>p«iL 



tion of petiole and blade, antl the stalked glands thickly beset 
tUe whole upper siirfnce of the latter. A sniiill insect aligUUiig 
thereon is helpless, and is soon touched by all the glands within 
reacliing distance ; also the blade itself commonly incurves, 
taking part in the general movement. It has recently tiecn 
(lemon Btrateil that the captureii insect is fed uixin, 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 chlorophyll), hare I 

also the power and the habit of obtaining ready-oi^nized food 
by enpturc, and are benefited by it. 

224. S]>ecies of Drosera inhabit most parts of the world, and 
the genns in numerous in species. A near relative, Diona-a, is 
of ft single species. D. museipula (\'enus'8 Fly-trap), inhabiting 
only a limited district in the sandy eastern Iwrder of North 
Carolina. It is more strikingly sensitive and equally carnivo- 
rous, bat 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 Uiis 

ra cloiaL Prulmblr ■ fl; !■ t 



be taken for the leaf-blaclo, the part below would be a broadly- 
winged foliaceous i>etiole. If the latter be the true blade, the 
apparatus in question must be reckoned as a peculiar terminal 
ap|>endagc. Both ait: modciatelj' green, and aet as foliage. 
Tlie s|>et'ially endowed temiinal |»ortion acta also in a deei<ledly 
animal-like manner. When cither of tlie three or four slender 
bristles of the ii|)i>er surface are touched, the trap 6iid<lenly 
closes, by a movement ordinarily quick enough to enclose and 
retain a fly or other small insect. The intci-erosstug of the stout 
mat^inat bristles detains the captive, unless it hap|)oiis to be 
small enough to escaj>e by the intenening little oi)eniugs. 
Otherwise, the siiles soon flatten and are brought flrmly into 
contact, and a glairj- secretion is [KHircfl out from numerous 
immersed glands : this, with the extracted juices of the mac-crated 
insect, is alter some time reabsorbed ; the trap, if in a healthy 
condition, now re-oiiena and is ready for another capture. For 
references to the now copious literature of tliis whole subject, and 
for its physiological treatment, the succeeding volume should I<e 

225. Leayes for Stom^. Xutritive matter is stored in leaves 
in many cases, aud not rarely in lea\'es which at tlio 
same time arc subseniiig the purpose of foliage. 
This occurs in all fleshy leaves, to a greater or less I 
extent, according to the degree of tliickcning or i 
accumulation. The leaves of the Ccntiuy Plant \ 
or Agave, for instance, are green and foliaecoualy 
elticient at the surface, while the wliole interior is 
a store-house of farinaceous and other mitiitious 
matter, as much so as is a |)otato. The leaves 
of various si>eeie3 of Aloe, Mesenihryanthemnm, 
Sedum, aud other "succulent" plants (in which 
a large part of the accumnlation is water) are not 
rarely so obese as to lose or much disguise the 
foliaceous api»enrancc. Sometimes one jwrtion of 
a leaf is of normal texture and use, wliile another , A 

is used as a re9e^^■oir for the nourishment which the r upi ■ H|Ui 
foliaceous part has produced. Fig. 232, a loaf from ^'- 1 - ."^J ^ 
the bulb of White Lily, the base of which forms « 

one of the bulb-scales, is an instance of the kind.' The most 

' In Dicenlra Cucullftrin ani! (more atrikinftly [mm tlie «pnraenM« of tlip 
grains) in D. Canadensis, the malltr clnborRleiJ in the much dissocted bindc is 
conTcjcd to tlic veiy base of (lie lonii; petiuli', and tliorc drpoeited in n con- 



decisive instance of lonves need for sloroge of food is i 
material proviaion for tlie nunrishnient of llie eralirjo in geritii- 
nation, in whicli the first learea, the eotyledoiis. are tunnel to tliis 
account. (21-37. Ac.) Alltr or 
wLilc disfliargiug this si>ectal diitj-, , 
tlic eot.vlwions muy fulfil their gen- 
eral office, by scniiig as foliage (as 
in Maples, Fig. 8. and Pumpkins, . 
Fig. 47) ; or. tlirotigli various intcr- 
tnctliate conditions, tbcy may be < 
wholly devot«l to Btorngc, -as in 
^Ihc I'ea, Oak, Uorseeliestnut, &c. 
(Fig. 37-13.) 

226. Leaves as Bnlb-BcateB, how- 
ever, are for the most part wholly 
applied to tliis use, being leaves 
re<iueetl to abort scales or to 

^coneentrif coats, and thickened 
throiiglioiit by nutritive deposit. 
The accnm Illation of such leaves 
forms the mass of the bulb, as of 
the Lily, Fig. 118, Onion, Fig. 
113, &c., alsoof bulbli'ts. (120.) 

227. LesTeH as Bnd-wales. being 
for protection of nascent parts, have 
been explained under buds. (70.) 
The evidence of foliar natuie af- 
forded by transition is well exhib- 
ited by the Sweet Buckeye, altliougli 
the n-liolcscries of gradations, from 
biid-scales to compound leaves < 
is seldom seen united in one Imd, 
as in Fig. 233. In this ease, the 
bud-Bcalcs are homologous with 
petioles. In Magnolia, tliey consist 

m of stipules (Fig. 81. 82) : in the 

LJIae, Uiey are liomologous with leaf-lilafles. The two pairs of 
l>nd-3CjileH wliich subtend and proleet through winter the nascent 
head of flowers of Cornus flori<la are morphologitally the apex of 

cciitmtpd condition. In Ihe (nrm of n solid gnin, wliiclt rcmaini for next 
ychT'i luc. Ihc wholp leaf pxctrpl ihii Ihickenpil bn«e dying nwoy al llie cloae 
of thp short ic&iion'g pviirtli. 

no Sn. LMtu Af k •laTvl'>|iliiE bu'l oT Uio I/iit Sw»I BiickcJV (^l>riiln> luinl- 
Sonl BhowlnE a ncati; cani|4ete Kt uT grwIsUuni from a xailc to a miup'iniiil loaf of 


blades. When the bloasoma develop in epring, these scales grow 
from beneath, greatly expand, and become obovate ot obcordate 
petaloid ka\ea, the brown terminal iiotcL 
of wliit'h is the bu<l-sciilc, wliich was un- 
able to take part in the vernal growth. 

227'. Leaves as Spines. All gradations 
may be found between spiny-toot lied Icai'es 
(as in Holly), in wliieh teeth are {Minted and 
indurated, and leaves which arc completely 
contractual into a simple or mnltiplc spine. 
Indeed, sucli a transition is seen in the Bar- 
berry, Fig. 234. The foliar nature of such 
spines is manifest from their ixjsition, suti- 
tcnding a bud from which the foliage of the 
season procectls, and themselves not sub- 
tended l>y any organ. In some Astragali, 
the i)ctiolc of a pinnate leaf inilurates into 
a slender spine and persists, tlic leaflets 
early falling. The spine in Fouqniera is 
portion of tlic lower side of the iX'tiole o 
midrib, indurated an<I persistent, the rest 
of the leaf sejiarating by splitting when it 
has sencd its office. ' ™ 

228. Leaves adapted to CliDiblnir. Some [ilants climb by the 
action of the stem or of certain branches si>ecially adapted to 
this pnqMse (00) : otiici-s gain tlie needful 
sup'port by means of their leaves (101) ; some- 
times lij" an incurvation of the tijra, either of 
a simple blade as in Gloriosa, or small partial 
tilades, as iu Adiumia, and often in Clematis, 
tliereby gm[){>liiig the support ; sometimes by 
tlic petiole making a turn or two aionud a 
supiwrt (as in Jlawrautlia, climbing Antirrlii- 
niima, Rliodoeliiton, and .Solanum jasminoitles, L 
Fig. 23r>) ; sometimes hy the transformation L 
of one or more leaflets of a comiwund leaf I 
into tendrils, as in the Pea and Vetoli (Fig. 
204) ; sometimes by the suppression of ail the 
leaflets and the conversion of tlie whole i>ctiole into a tendril, as 
in Lathyrus Aphaea (Fig. 219) ; and perhaps by the con\crHion 

1 Dotcribcil in PUntte WriplitiniiH!, ii. C3. 
Flo. 23t. A venial Kliont orioium™ Burberry, i-liowlng a lower l™r In tlio ii.irniiil 
•Ula; IhanaiCnartlKlly. thoHMllIhle>ier«iu|il«lelr, Ininsr<>rniallnti>i>r>liiM. 

F10.!3S. SaluiiimJ»HDlnDld«,cl]mt)lD|[b]rcuUlnguiiMliiiigtblnilnratiiigiici<Dl». 



of a pair of stipules into tentliib in Sinilax. At least the t£ii- 
drila htre occupj- the [wsition of adnuti; stipules. Tlie tuiirtrila 
of CueiubitacciB are peciiliui' uuci amiiiguoiis, on account of tlii'ir 
lateral anil cxtra-axillaiT |>oBition and the manner in which the 
(.vmjxiund ones develop tlieir bi'anehca. But they art; doubtless 
pailij- if not wholly foliar.' 

^21). PetaloU Leaves, Bracts. Certain leaves, situated near to 
flowers, and ilevelopiug little or no chloi-opliyll in their paren- 
cl>.\'ma, exc-hanjic llie oiiUnarj- green hue and herbaceous textui'c 
for the brighter colors and more delicate Btructiire which are 
commonly seen in and tlionght to characterize Ho wei'- leaves. 
Sucli arc said to be colored, moaning, od applied to foliage, of 
some otlicr color than green. As petals ore the lv{)c of such 
eoloi-ed paits, they are said to be petalotd, i, e. |>ctal-likG. They 
are like jKtals, moivover, in one of the purposes which tliese Bub- 
8er\'e. (293.) Examples of these petaloid leaves are seen in the 
shinbby Mexican Euphorbia called Puinsettia, in Salvia splen- 
deiis, most sgwciea of Castilleia or Fainted Cup, also in tlic 
wUitc hoo<l of Calla and Ricbardia JEtliiopica (called Calla Lily), 
and in tlic four wliitc leaves which subtend the flower-head of 
CornuB floridu, anil of the low lierlmeeous Cornel, C. Canaden- 
sis. (Fig. 294.) Sucti leaves, being in proximity to flowers, and 
all others which arc within a flower-cluster or aiv borne by 
flower-stalks, I'ceeivc the sjiecial name of Bracts. More usually 
bmets are not pctaluiil, l)nt diflcrent in size or Blia|je from ordi- 
naiy leaves, eitlicr l)y abiiipt cliange or gradual transition. Not 
uncommonly tliey are reiliiced to scales or mere rudiments' or 
vestiges of leaves, of no functional importance. 

S3<J. Flower- LoavM. Tlie niorpholog}- of Ictnes extends not 
only to "the leaves of tlie blossom," more or less accounted as 
such in common parlance, but also to its peculiar and essential 
ui'guns, the relation of wliieti to leaves is more recondite. Their 
nioqihology neeils to be treated scparatflj-. anil to Ik preceded 
by a study of the aiTangemcnt of leaves and of blossoms. 

' Till! niDHi Btttlsfuctory initTpretfttion may \k llint of Brnun ami Wyillw, 
Hil..pteil by Eidilcr (BUillitnilingnimme, i. 304); tlmt tin- rtowi-r of Cucur- 
bita and iti pvilunol? reprctcnl Ilii^ axillary brant'li, ilic tonilril I17 it* slile 
aniwcrs to one of the bnu;ileti [tliat of tlie other shiv W-\i\g >upi>rcMi'il), 
anil the aupemuineniry branuh iprings from the axil uf llie tendril. Tliii 
■nakea of the tendril ti liniple leaf, of which (he bntnehca nrc the rib*. But 
the tenilril-divisitina are vvidenily ilevelopeil In spiral oriUT. and in vlgonms 
gmwtlu nceupy different lidghli on the lendril-nxi*. Thii favor* Kaudin'a 
view, that the main tendril U cauUoe, anJ iu Uivitiont leavei. 




Section 1 

The Distbibltios of Leaves on the Steu. 

231. PiivLLOTAXv (or Pli^loiaxis) is the stuily of the distiU 

biition of leaves u|M)n the Btem and of the laws which govern it. 

The general euiicUisioii readied is, that loaves 

are distributed in a manner to eeonomize 3|ince 

and have a goo<l exposure to light, &c., and 

that this economy on the whole results from 

the formation of leaves in the bud over the 

widest interxals between the 

leaves next below.' Leaves 

are arranged in a consider- 
able variety of ways, wliieb 

all fall under two modes, tlic . 

VeTticilUiU and the AUer- 

nale (13), but whk-h may 

also be termed the Cyeliccd 

and the Spir<d. 
•2Zi. Alternate leaves are 

those wliieh stand singly, 

one after another; that is, 

with one leaf to eneh node 
or borne 

on one height of stem. Vertieillatc leaves are 
those with two or more at the same height of 
stem, dreularly en(.-om|iassing it, t. e. fonning 
a Vtrtietl OT Whorl. Vertieillate and whorle<l 
arc synonymous tenns to denote this arrange- 
ment. Tliese two kinds of leaf-arrangement 
are commonly rankctl ns three, viz. alternate, 
op|)osUc, and whorle<l. But tlie opiiosite is 
only the simi>lest ease of the whoile<l, lieing 

' Fur the moal cnrnprcliL-naivc iliscusaion of phyllotaxy in cinincclinD with 
ilpvelnpmcnt, &nil in view nf theav rclotinnB, ki' Hnfrnviflpr. Allfcemrine 
Morphologic, % II, «nd Ch«uncey Wrighl, Mem. Amer. Acadcniy, ix. 389. 
FIO. 330, Alternate, m, Oppaette, 238, Vetlk1ll>M or wborled leares. 



Uial in which the members are i'ei1iice<l to two. This case 
BO much commoner than whorla of three and. ol' higher nitmljera 
that it took fVom the first its special name uf opposite, so thnt iu 
(lescriptions tiie phrase " leaves veiticillntc " implies more tlian 
two leaves in the n-Iiort. But it sliuutd t>c kept in miud that 
"leaves opposite " is the same ns " leaves in whorls of two." 

233. The greater number of phienogamous plants (all but the 
monocotylwlonons class) begin with verticillate leaves, mostly 
of the simplest kind (t. e, cotyledons opiiosite) : some contuiue 
vcrticilUHe throughout ; some change in the lli'sl leaves of the 
plumule or after the first pair into alternate, and again into 
verticillate in or toward the blossom, in the interior of wliicb the 
alternate ariiuigcment may be again resumed. As Xalure jiasses 
readily from the one mode to the other on the same axis, we 
may ex|>ect that the two may be comprised mider some common 
expression. But they have not yet been combined, except by 
gratuitous or somewhat foi-ced IiyjKithescs ; so that lor the 
present they should be treated in morphologj- as primarily dis- 
tinct arrangements.' 

234. Verticillate or CrcUcal Arrangement. Ilci-e the leaves 
occupy a succession of ch-cles, or form whorls around the stem, 
two. tln'ee, four, five, Sic, in each whorl. Aei-ording to the 
uiunlrer, the leaves are oppotice, lemale, qtialemit/e, tjuinnie, unci 
so on. The characteristic of the individual whorl is that the 
memlicrs stand as far apart from each other as their nimiber 
renders possible, i. c. they divide the circle etjiially. Thus, when 
only two, or op|>osite, their midrilis or axes of insertion have an 
angiihir diveigence (as it is termed) of 180°; when three, of 
120° ; when four, 90" ; when five, 72". 

233. The characteristic of the whorls m ■'elation to each other 
is, that the members of successive wliorls stand over or under 
the intenats of the adjacent ones. In other wonls, successive 
whorls allrrnate or rffcutsate. This eeonomiites simco and light, 
or gives the liest distribution which the cyclical system is ca[w- 
blo of. And-it is in acoonlanee witli tlie general conchision of 
Honnei9tei''s investigation of the origin of phyllotaxie oiTange- 
ments iit tlio nascent bud, vi2. tlial new members originate just 
over the widest intcn*als between their predecessors next below. 
Tlitis, in opijosite lea\ea or wliorls of two (I'"ig- 237), the suc- 
cessive pairs dreu*saie or cross at right angles, and so four 

' It u n-ailily vi'ii tlial wliurla i<i«y be pruduccil by tlm iiiin-i]<>vvlu|iiiient 
uf llie inli-mudi'* Iielwivn llie Icbvi'b uf a acriM of two. thm-, fivi>, or niurv 
in Nllcnialc unlpr. TUt djffli'ulty u lliit Ihe membpre of thv next whorl do 
not fcillow the order that tlicy shoulU upon tlili BtippoiitioD. 



straight equidistant vertical ranks are produced. In temate or 
triraerous whorls there are six vertical ranks ; in quatemate or 
tetramerous whorls, eight vertical ranks, and so on.* 

236. The cases in which successive © 
paira of leaves do not decussate at right 
angles, or the members of whorls are not 
exactly sui)eri>osed to inten'als, but as 
were wind spirally (as in Dipsacus, many 
Car3*ophyllaceffi, &c.) , may some of them 
Ikj explained by torsion of the stem, 
such as is very manifest in numerous in- 
stances ; and others may be ixjsolved into 
instances of alternate leaves simulating or passing into whorls 
by the non-development of intemodes.^ 

237. Alternate or Spiral Arrangement. Here the leaves are 
distributed singly' at different heights of the stem, and at equal 
intervals asresi)ects angular divergence. (Fig. 23G.) This angu- 
lar divergence (i*. c. the angular distance of an}' two successive 
leaves) differs in the various kinds of this sj'stem of phj'llotax}', 
but is always lai-gc enough to place the leaves which immediately 

* These vertical ranks have, by some German botanists, been named 
Orthostichies ; but this teclmical Greek is no clearer and no shorter than the 
equivalent English, which answers every purpose. 

* In Liiium 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 nonnai 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)A *"^ (i)A- 

Braun's mode of notation for the onlinary succession («. 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 i 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 also some cases of intermediate divergence; such as (|)| in the 
upjier leaves of Mercurialis perennis, (i)^ on certain stems of Linaria vul- 
garis, {\)-ff exceptionally in the leaves of Epilobium angustifolium and the 
scales of Norway Spruce, H)-^ exceptionally in the scales of Norway 
Spruce. See Ordnung des Schuppen an der Tannenzapfen, 376, &c. 

FIO. 239. Groand-plan diagram of six trimerous whorls, showing their alternation. 



follow each other in the ascending order npon different sides of 
the axis : it also secures an advantageous sjmeing of the leaves 
over the whole length of tlie axis. Their vertical distance from 
each other of course dei>cnds on the length which the intemodca 
attain, which is a niatt«r of growth and is very variable ; but 
their angular distance is Hxed in the kind or numerical plan of 
the particular phjllotaxy, an<l is uniform throughout. 

239. The leaves arc said to be allentale, because they come 
one after another, uow on this side, then on that, as they ascend 
the stem. The arrangement is said to be tpirtd, because if a line 
be drawn or a thread e\-t«ndcd from the base or insertion of one 
leaf to that of the next higher, and so on, taking iu all the leaves, 
it forms a helix, more or less loose or close according to the 
development of the intcmodes. (See Fig, 242.) This imagined 
spiral hue nawnds cootiuuously, without a break ; and on it the 
leaves are equably laid down.' 

'lA'i. Almost all the ordiuaiy instances of spiral phyllotaxy 
belong to one series, having very simple arithmetical relations. 
So that this may !« taken as the type, and the few others re- 
garded as exceptions or somotimes as modilications of it. The 
kinds are simply designated by the number of vertical ranks of 
leaves : they are tcclinicoUy named by prefixing the proper 
Greek numeral to the word meaning row or rank. The arrange- 
ment called 

Dittiehom, or Two-mnltd, is the siniplest and among the com- 
monest, occunniig, as it does, in all Grasses and many other 
mouocotj'lcdonous plants, in Lindens, Elms, and many dioo- 
tylo<Ionoua genera. Here tlie leaves are dist>osed aitematelj' on 
exactly opposite sides of the stem (as in Fig. 1) ; the second 
leaf being the farthest imssible from the first, as is the third from 
the second ; tlie third tliereforc over the first, and the fourth over 
the second, and so on, thus forming two vertical ranks. The 
angular divei^nce is here half the cireumference. or 180* ; and 
the phyllotaxy may be rei)resented hy the fVaetion i, which desig* 
nates the angidar divergence, while its denomiuator expresses 
the number of vertical ranks formed. 

Triakhovt, or Three-rariixd, is the next iu the series, and ia 

' But when »p ronch a le«f which ttanAt diroclly over a lowpr ami older 
one. we my that one sel or tpire \i eoniplftpd, and that tlii» leaf ia ihe flril 
of B succpeding sel of »)iiiy. From nnalogy of «ucli an open ipin to the 
clovird t'j-i'li; of a whorl nt IvftTf*. it li nol unuanul to iletiKiiwIe tl 
likewise aa k cydr. Yet It ii better ( with Eichter) to restrict that tenn. and 
the adjpoiive egtiical, to Tcrllcillale phyllotaxj, or to whurli, to wbkh it 
proiwrly and etymological!/ Iwlonipj. 



less common, though not rarc in monocotyledonous 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 J designates this 

arrangement. The angular r^sai « e 

divergence, or distance of 
the axis of the first leaf 
from the sec*ond, and so on, 
is one thiitl of the circum- 
ference (or 120°) : conse- 
quentl}' the fourth leaf comes 
over the first, the fifth over 
the second, the sixth over 
the thiixl, 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.* 

Pentastichous, or Five-rankedy sometimes termed the quincuncial 
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 angukr distance from thp firat to the 
8:^cond leaf (passing the shorter way) is J of the circumference, 
or 144°. But tlie spiral line makes two turns round the stem, 
on which six leaves are laid down, with angular divergence of §, 

* 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 utalk, with the sheathing bases of the leaves, of a Sedge-OraM 
(Carex cnis-corvi), showing the three-ranke<I arrangement 241. Diagram of the crow- 
section of the same. Tlie leaves are numlwrwl in succession. 

FIO. 242. IHafn^ni of position of nix leaves in the fl ve-ranketl arrangement : a spiral 
line is drawn ascending the stem and passing througli the successive scars which mark 
the position of the leaves from 1 to 6. It is made a dotteil lino where it passes on the 
opposite side of the stem, and the scars 2 and 5, wlilch fall on that side, are made 
fUnter. 24.'J. 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 
torn; at the sixth would come directly before, or within, the first. 




and the sixtli is the first to como ovpr any one below; the 
seventh comes over the second, the eighth over the tliird, &c. 
The Icavt's are thus brought into flvo vertical ranks ; but those 

§11 five loaves are laid clown on two turns of tho 
'° , • helix (the sixth Iwginning tlie aeconil revolu- 
g ' tion) ; tlie angular divergence of the leaves in 
' , • oriler is J, or H4° : the angular distance of 
1 ' the vertical ranks, li". This is a vcrj- advan* 
•M tageous disti-ibiition for ordiuury foliage on 

erect or ascending branches. Its formula is j(, oxpresaing the 
angular divergence, the denominator also indicating the number 
of vertical ranks, the nun 
ator indicaliiig the number > 
of revolutions made in add- |^ 
ing one leaf to each rank, i 
Fig. 344 illustrates this ar- 
rangement on a cone of 
American Larch, the scales 
of which are homologous 
with leaves, the numl>ora 
in sight are alHscd. and those of the whole 
cone displayed on a plane at the side, 

Ociostickout. or Eight-ranked, a less common 
arrangement, occurs in the Holly, Aconite, 
the radical leaves of Plantago. It has the 
angular divci^^nce of 135°, or \ of the cir- 
cumference, and the leaves in eight ranks, 
tlie ninth over the first and at tho completion 
of tho third revolution : it is therefore repr&. 
sented by the fraction |. 

240. The obvious relations of the fractions 
J- !■ ?• t> representing tho primary forms of 
B|jiral phyllotaxy, ore that the sum of any two 
lumerat^n-s is the numerator of tlie nest suc- 
ccc<ling fraction, and the same is true of the 
deuomiuators ; also the numerator is the same 
as the denominator of the ni-sl but one pre- 
ceeding fVaction. Following Uiese indications, 
the series maybe extended to ^, /,, JJ. %\, &c. Now these 

no. iU, A aniB of Ui« •nuU-rrulud Americmi Lorcti (Larii A 
as Kftlis uumtwtul. ollittilUiig Ilia Uic-nuiknl ■rntngEnivnt. 
• Fit). 3U. Anntttet nr Ihe Hoiue1a>k.r(lilbhlnelbDS-l3>unn(ui 
In sljEbt iimulHiraJ. Ih« l»li uver Uia ftiU. Ilic IMbovsr theotb. Sr 

Fit)- 3W. Kmyont Willie PinoiriDiuStniliiuJwIUinulaiiuiubci 
viil Futuu Kvoiubftry jf|UiuLii Imukud. 


cases actually occur, and ordinarily onl}' these. ^ The -^f^ and 
/j are not uncommon in foliage. The rosettes of the House- 
leek exhibit the i'*jj or thirteen-ranked arrangement, as also does 
tlie cone of Pinus Strobus, the 14th leaf falling over the first. 
(Fig. 246.) The ^ 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 \% 
and Ji phyllotax}' may be readily made out. This actual series, 
^, J, §, g, &c., answers to and may be expressed by the con- 

tinued fraction, 4 -f- j. 

^ "*■ + + !, &c.' 

1 When other instances are detected, thej are found to belong to other 
series, following the same law, such as the rare one of \i \y f , -^. 

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 docs, in fact, possess in a higher deg^ree 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 toW* * 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, which 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, acconling 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 
f and ^t -fi, &c., which are all indistinguishable as measured values in the 
plant, do actually realize this property with all needful accuracy. Thus, 
I = 0.C26, ^6, = 0.615, and y^ = 0.619, and differ from k [the ultimate value 
to which the fractions of this series approximate, or what is supposed to be 
the tyx)e-form of them] by 0.007, 0.003, and 0.001 respectively ; or they all 
differ by inappreciable values from the quantity which might therefore he 
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." — 
Chauncey Wright, in Mem. Amer. Acad, ix. 387-390. 



241. The four leading gro<i<?s of altemat* loaf-arraDgemont, 
namely, those with angular divergence of 

i = 180° I = IW" 

i=120'' i = 135' 

are represented by rational fVactions, or have the angle of 
divergence commensurate with the circumference. The leaves 
should therefore be in strict vertical ranks, or, to use the terra 
proposed by Bravais, reetherial. But bejond this grade the 
angle of divei^nee hecomesirrational to the circumference (in the 
yi, = 138" 27' 41.54", in jPi = 137" 8' 34.29", in the deduced 
typical angle to which these higher forms more and more 
appro:£imate. 137° 30' 28"). and so theoretically no leaf ia 
exactly superposed to any prece<ling one. The leaves thus 
distributed on &□ inSnite cun'e are by Bravais said to be 
ctini*eriat. But (as stated in accompanying note) the devia- 
tion from verticality is practically inappreciable, and even the 
dilferetice between ^ and }}, &q., is too slight and variable in 
many cases for certain determination. 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 leases would be dis- 
tributed most thoroughly and rapidly around the 8t«m, exposed 
moat completely to light and air, and provided with the greatest 
freedom for sj-mmctrical ejqiansion, together with a compact 
arrangement in the bud." Even in the simpler grades of com- 
monest occurrence, each leaf (accortling 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 ia 
com|>lcted, when the new leaf is placed over an old one.* 

242. It is to Ije noted that the distichous or I varietj- gives 
the maximum divergence, viz. 180°, and that the tristichous or 
4 gives the least, or 1 20° ; that of the pentaslichoiis or g is nearly 
the mean between the first two ; that of the J, nearly the mean 
between the two preceding, &c. The ilisadvantage of the two- 
ranked arrangement is that the leaves are soon superjiosed and 
so overshadow each other. This is commonly obviated hy the 
length of the intemodes, which is apt to be much greater in this 
than in the more complex arrangements, thcaToro plac" 
vertically faithcr apart ; or eke. as in Elms, Beeches, 

I the 

' Tliii L-om-spoiiil* with Ilofmtwlcr'e g;cncT>t nile, that " nev Utrrnl 
iin'mlMT» hare their orifrin ivbjivc llie widest gsps belwetn the inecrlions of 
llie nearcil older mctnbcn." Yel the fntt Ilint tlie clmrButer of llie le«f- 
nrranft^mcnt U laid down at the Ix'ginning In Ihe bud doe* nui go tnr id llie 
»ay ut tlie mecbBnii;al rxplnnallon wlili-'h he hivukts. 


like, the branehlets take a horizontal position and the peti- 
oles a quailer twist, which gives full exposure of the upi)er 
face of all the leaves to the light. The J and §, with dimin- 
ished divergence, increase the number of mnks ; the J and all 
beyond, with mean divei*gence of successive leaves, effect a more 
thorough distribution, but with less and less angular distance 
between the vertical mnks. 

242*. The helix or j^rimitive spiral upon which the leaves 
successive!}' originate ascends, sometimes from left to right, 
sometimes from right to lefl,^ commonly without change on the 
same axis, and prevailingly uniform in the same species ; but 
occasionall}' both directions occur in the same individual. The 
earliest leaves of a stem or branch, or the last, are often on a 
different onler from the rest ; or (as already stated) the spiral 
ma}' change into the cyclical, or vice versa. 

243. The relation of the ph^llotaxy of a branch to the leaf 
from the axil of which the branch springs is somewhat various. 
But in Dicotyledons, the firet leaf or the first pair of the branch is 
mostl}' 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 usuall}' 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 b}' torsion 
of the axis : indeed, this maj' equally occur in short internodes, 
sometimes irregularl}' 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. Tlie 
later leaves of Baptisia ixirfoliata, which are nonnall}' distichous, 
bec*ome one-ranked by an alternate twist, right and lefl, 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 
tliese vertical ranks are commonlv 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 b}' some parcutichies. These oblique spiral 

^ That is, of the obsenrer and as seen from without. See p. 61, foot-note. 


ninks ai'e a Hcceseary consequence of tlio regular asccniliag 
aiTangoniPnt of pni'ts witb e(]iial intervals over the drcumtbrcncc 
of tbc axis ; and, it' tiio Icaius am □umb^i'ed ooiiseoutivclj', tbeir 
numbers will iietessarily stand in ailtbinetical pi'ogrcssion on the 
oblique ranks, antl have certain obvious relations with tlie pri- 
mary spiral which originates tbem, as will he seen by projecting 
them on a vertical plane. 

24o*. Take, for example, the } arrangemeilt, where, as in the 
dii^Bm anntixeil to Fig. 244, the primitive spiral, written on a 
plane surface, appears in the nmnbers, 1, 2. 3, 4, d, C, and so 
on ; the vertical ranks thus formed are necessarily the nnmliers 
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. l-^-."), which, 
if we coil the diagram, will be continued inlo 7-D-U-13-15; and 
also the 2— 1-C-H-lO continues into 12-14, and so on, if the oxia 
be prolonged. Here the eircumferenc* is occupied by two seuon- 
dary left-hand series, and we notice that the comniini rliflcrence 
in the sequence of numbers is two ; that is, tbc number of the 
parallel secondarj' spirals is the same as the common dilfercnce of 
the numbers on the leaves that compose them- Again, there are 
other parallel secondaiy spiral ranks, three in tmmber, which 
ascend to the right ; viz. 1-1-7, continued into 10-13; 3-6-D-12, 
continued into 15; ond 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 proi)er numbers on the leaves, 
when they are too crowded for directly following their sncces- 
8ion, and thus to ascertain the order of the primary spiral series 
hy noticing what numl>ers eomc to bo 8U])erposeil in the verti- 
cal ranks. Thus, in the small cone of the American Lnrcfa 
(Fig. 244) , which iisnally corajilel^s only tbi-oe heights of leaves, 
llie lowest, highest, and a middle one make a vertical row 
which faces tlie oiisencr. Marking tliia first sc-ale I. and count- 
ing the pamllel secondarj' spirals that wind to the left, we find 
thiit two occupy the whole circumferent*. From 1. we number 
on the scales of that s|)iral 3-5-7, and so on, adding the conf 
mon ditferencc 2, at each ste|i. Again, counting from the base 
the right-hand secondar>- spirals, we find three of them, and 
therefore proceed to number the lowest one liy adding tliis oom- 
mon difference, viz. 1-4-7-10; then, ]>assing to the next, oa 
which the No. 3 has already been fixed, wo carry on that se- 
quence, 6-9. &c, : and on the third, where No. 5 is olr*-ady 
fixed, we continue the numl>ering. R-ll, &c. This gives us in 
the vertical rank to which No. 1 Ix'longs the Bcqncncc l-6-ll, 
showing that the pbyllotaxy is of the five-ranked, or J order. 


It is ftirther noticeable that the smaller number of parallel sec- 
ondary' spirals, 2, agi-ees with the numerator of the fraction in 
this the J arrangement ; and that this numl)er, added to that of 
the parallel secondary spirals which wind in the opi)osite dircction, 
viz. 3, gives the denominator of the fraction. This holds good 
throughout ; so that we have only to count the numlxjr 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 numlwr 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 onl}' the 
most manifest secondary spiral of each direction in oixler to 
lay down the proi)er number on the leaves or scales, and so deter- 
mine the phyllotaxy.^ In a rosette of the leaves of Ilouseleek 
(Fig. 245) and a cone of Pinus Strobus (Fig. 24G), the num- 
bers which can be seen at one view are apjxjnded, 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 sui)eq)osed to the first, indicating the 
^ 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 ma}' suffice for 
botanical uses. 

24G. Relations of Wborls to Spirals. Verticillate and alteiiiate 
ph^'llotaxy, or whorls and spirals, in all complete exemplifica- 
tions, are to Ik? considered morphologically as distinct modes, 
not to \}c practically homologized into one. Nevertheless, transi- 
tions between the two, and abrupt changes from one to the other 
on the same axis are not uncommon, the former esix?cially in 
the foliage, the latter in the blossom. If the spiral be assumed 
as the fundamental onler, it is not difficult to form a clear con- 
ception as to how such changes come to pass. A single whorl 

^ In applying this method to the determination of tlie pliyllotaxy of a 
cone, or any such aasemblage 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 ))lninly mack' up of pairs of oppo- 
•ite scales which are spirally arranged. See note under 2^)0. 



may most naturally be produced by the non-developraent of the 
iiitcniodes bctwuen any two, three, ov more altJ^riiate leaves. 
Two proxiniat* distichously alternate leaves would thus form a 
jiair ; thii thi'ee leaves belonging to one turn of the spiral in the 
tristU'hous (J) arrangement would oomiKise a trimeroua whorl ; 
the Bve leaves of the two turns in the pentastiehous (f ) arrange- 
ment, a S-merons whorl, &c. Veriticatioiie of this conception, by 
whorls breaking up or reverting to spii-ab, are occasionally met 
with, and tlie successive overlapping in spiral order of the 
meml)er8 of a trimerous or pentamcrous whorl is very common. 
The few instances among phrent^amous plants in which the 
leaves are opposite and all in the same plane ' (that is. the suc- 
cessive pairs supeqwsed) may be deduced from the ilistiehous 
alternate mode becoming opposite without furtber change, by 
the simple suppression of alternate intemodes. The frequent 
disjunction of the members of the pair in similar and analogous 
cases goes to confirm this view. But the characteristic of whorla 
ordinarily is that proximate whorls alternate, that pairs de- 
cussate. We cannot boraologtze this with spiral phyllotaxy; 
for in this lies the fundamental difference between the two plans. 
We can explain it only by a reference to Hoftneister'a law, whieli 
generally governs leaf-origination as to position, namely, that 
succeeding leaves appear directly almve the intervals t>etweon 
the nearest preceding (241, note): this gives decussation or 
alternation of successive pairs or whorls.' 

247. Hjpothesh of tho origin of both. Instead of regarding 
the spiral path on the stera which conueuts successive alternate 
leaves as a pnivly formal representation, it may t>e conceivetl to 
be the line along which the members in some original form were 
phj-sicalty connected, in the manner of a leaf-like cx|>ansioti 

» At in Loranlhaa EuropisuB. *c., seenrding lo Braun. See 230, note. 

* Thla rendtre the Tertit-'illsle nn ailvuntoftvous amuigenii^nt, perliapa no 
leii so than Ilie diittribuliun which «pinl pliyJlolnxy cQei-'U, Both miut be 
considered to have been delermined by and for Iheir respective utilitie*,and 
to hare been independent dclerminationi. For " Ibcre it no eonlintutj' or 
piindple of connc<:Iioii between spiral srrnngemenls and whorli " (Cliaunce^ 
Wright) i (ince, although indiildual whorls are edsily rfducible to spiral*, 
each iDceesaioo is aa absolute break of (hat system. 

As whorls of four memben often (ai especially in calyx. bracts, i,e.) laty 
and sometinien should be riowed a* two apprnxlmatc pairs, so even ihc tplrsl 
of fl*e members, ni in a qufticunciai calyx, has been conceived lo coniiit of 
two whorls, one of two, the other of three leaves, the second alternating wllk 
the flrtt aa nearly as ponible. Bat ihit appears far>fetched and o{ loose 
appUcatiun. It is much clearer as well as simpler to regard the altemalo 
as the f nndameiiCal phyllotHxy, and to deduce indiiidoal whorls from spirals, 
if need be, rather than to iiuapine ipimli as somehow evolTed from whoil*. 



resembling a spiral stairway. Upon this snppoaition, the leaves 
would be tlio i-elics, or rather tlie advantageous results, of the 
segmentation ot such a frond-like 
expansion, the segments scjtaratcd 
through the development of the 
stem in length and flrmncss, and 
modified in the various adaptations 
to the conditions of higher vege- 
table life ; even as leaves themselves 
are modi Bed into tendrils, bud- 
scales, petals, or other usefully specialized structures. The tj-pe 
on this conception would be a ih>nd, consisting of 
an elongating axis with a continuous leaf-blade on 
one side, and this taking a spirally twisted form. 
But the fVond of Fucaceous Algee, Hcpaticse, 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 vertieillate arrangement.' 

248. Fascicled LeaTCS need to be mentioned here, 
in order that they may be excluded from phyllotasy. 
They are simply a cluster or tufl of leaves, belonging 
to more than one node, and led in a crowded con- 
dition because the intemodes do not lengthen. They 
may belong either to the alternate or the vertieillate 
series. Jn Barberry and in the Larch (Fig. 247), 
they are evidently altemntc ; and they may be inferred 
to be so in Pines (Fig. 248) , or even may be seen 
be BO in the bud-scales which form the sheath e 
rounding the base of the 2, 3, or 5 foliage-leaves. 
In Junipers, the leaves of the fascicles are in the 
vertieillate order. 

I lliis ia the conception of the late Chauncey Wright. See hia elaborate 
and must suggeitive esMy in Mem. Amcr, Acsil, Arta and ScicnL-ce, acr, 2, 
li. 379. mainly reprinted in Phltoaophical Diacusaiona (poathumoua), 200-328, 
in whii^h tlie whole aubject of phyllotaiy is acutely discuwed, eapecially 
in Jls reUtioQ to queations of origin and developed utjlitiea. His conccpllon 

t^ ^IerTiiitolG4Taof thecleTcIoped ails of th« prec^lliig year Bepars1«<l. 

FIO. »8. PiMeat* bnncb of Pitch Pine, •ritb thne ■»•«• In a fMHrla orbnndle 
IntlMaillof achlD>calal<i)Hhich aiuwera Is a leaf of tbe main uli. Tbe bumlle li 
rarroandeil at tha b«« by a abort slieatli. formeil at the deUcale seals of Uie ailUarr 
bud, of whldi Uie three leaTea are tlis deielopeil (bllaca. 



Section II. Disposition of Leayes in the Bcd. 

249. Tematlon and fstlvatioo are terms in general use, under 
which the dis|>o8itioii of leaves in the bud is treated. The first 
relalPB U) oi'dinary leaves in this earlj- eonditiou ; the Bc<:oiid, to 
the parts of a flower-bud; not, however, as respeets iHierlion, 
or iKieitton on Uie axis, whkh is pliyllotaxy (231), but as lo 
j«, -50 (ji the ways in which they 

are coiled, folded, ovcr- 
lap|wd, &c.,either/iw«e 
orinterte. Prmfoliation 
and PrttfioTOtion aro 
ctymologieally better 
temis, substituted by 

250. The descriptive 
terms nliieh ivlate to 
individual leaves or 
>M parts, whether of foli- 

age or blossom, mostly range themselves under the lieada of 
plications or of enrolling, and are such as tJie following, the 
sectional diagrams of which are copied fW}m the original flgui-es 

would mnke the two plant equally primordial. But the freedom with 
wliich [liMc actually Intert^liange on the Mine axii greatly favore the Icm 
liypot helical licw that whorli may be condensed Kplrnls. Tliia acaumM 
only the well-known fact thai iniernndei may be completely non-tleveloped. 

' Better formed and more exprcwlve (erme: 1>ul the Linnsan one* are i 
tnoat Innie.and, though fuicirul, are not mi aleading. In EnKliili doauriplian, 
<t is u cDnrenieni and cqimily (ene to say that the parti are imbricate, ¥■!• i 
rat?, &c., "in the buil." Linni-iu. In tlic PliUosophia Botanic*, described ' 
these dispogitiona of leaves in the bud under tlie term Fel'ialio, — not ■ happy 
name, — but did not treat of them in the flower-hud. Later, in Temilui 
Botanic! (AmiEn. Acad. vi. 1702, reprinted by Gi»cke in IT81), he intro- 
duced the words Vrmatlo and jf.tllealK in their now current botanical aenic, 
to designate, riot (he time of lenfing and of flowering (spring and suin 
condition), bnt the disposiliiin of the parti in the leaf-bud «nd flower-bud . 
(at least of the petals) ai respecK foldings, coiling, &c., of single pans, and ' 
modes of overlapping or otherwise of contiguous parts. The terminology 
M regards single leaves, Unneus fixed nearly as it now remains. Th»t of | 
leaves or their homologuci in connection, and as respects the Sower-bud, waa 
very imperfectly developed until its importance (and much of lis tennl- 
nolofcy) was indicated liy Robert Brown, in his niemnir on Proteacee, 180B, 
hi the Prodromni a year later, and In other publications. 

Pii/Tii (the Grtvic name) is coming into use as a RcntTal term for Uie 
folding, lie., of single parts. 

FIO. m-!M. Llniiesn lUafreiiis of seeUans of learn In Ilwbud. 2m. CondulJI- 
nle. :;sn. FllnitE nr pWlad. 2SI. CanTOlnta. Va. Kefolutn. 3U. Invqlots. VM. 


in the Pbilosopbia Botanica of Linnieus. They were applied 
onl}' to foliage, but the}- are equall}* applicable to floral paits. 
Leaves, and all homologous or similar organs, if not simpl}- 
plane, will be either bent or folded or else more or less rolle<l up 
in the bud. The first thi-ee of the following tei-ms i-elate to the 
former, the remaining terms to the latter. They are as to the 
mode of packing 

Plicate or Platted (Fig. 250) , when folded on the several ribs, 
in the manner of a closed fan, as in Maple and Currant. This 
occura onl}- in certain palmatel}' veined leaves. 

Condupltcate (Fig. 249), when folde<l lengthwise, or doubled 
up flat on the midrib, as in Magnolia ; a ver}- common mode. 
The upper face of the leaf is always within. 

Reclinate or InjUxed^ when the upjKjr part is bent on tlie 
lower, or the blade on the i)etiole, as in the Tulii>-ti'ee (the blade 
of which is also conduplicate) . 

ConvoliUe (Fig. 251), when rolle<l up fix)m one mai'gin, i.e. 
one mai*gin within the coil, the other without, as in Apricot 
and ChciTV. 

Involute (Fig. 253), both margins rolled towaixl the midiib 
on the upiKjr face, as the leaves of Water Lily, Violet, &c. ; 
also the i)etals of Steironema and Tremandra. 

Revoluie (Fig. 252), similarly rolled backwaixl from both 
margins, as the leaves of Azalea and Rosemary. 

Circinal or Circinate (Fig. 254), when coiled from the aj^ex 
downwanl, as the leaves of Drosera and the fix)nds of all the 
true Ferns. 

Corrugate or Crumpled^ as the i^etals of a Poppy, applies 
to the irregular crampling of the otherwise plane comlla-leaves. 
Tills is a consequence of rapid growth in length and breadth 
in a confined space. 

251. The PtyxU (or folding, &c.) of an individual leaf, of 
which the foregoing mollifications 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, t. e, in aestivation. To this the 
following exposition is devoted, although sometimes applicable 
to leaf-buds alssp.^ 

252. The disposition of parts in sestivation, in respect to 
each other, is the result partly of their relative insertion, that is 

^ In the succeeding paragraphs, it becomes necessary to presuppose so 
much knowledge of tlie flower as is implied in tlie free use of such terms as 
calyx and corolla, sepals or calyx-leaves, and petals or coroUarler' 
if need be. Chapter VI. Sect I. 



Uieir pliyllotax}-, anil partly of the way in wLil-Ii they vom 
when tUeir margins meet in gronth. Tlioac lea\'e3 wliicli are 
witUin, or of liiglier insertion on tlic axis, will almost ueee»saril}- 
1m! enclosed or overlaiJi>e<l ; those wliieli ore inemlters strictly 
of the same whoil or i-ycle may (iiil to (»me into L-ontoit, or 
muy meet without overlapping at the contignous mai^ins or 
apex ; yet they may be on;rliip[>eil. since tliey may have grawn 
imeqnallj- or some a little earlier than tlitjir fellows. Conso- 
(jnt-ntly, no perfectly clear line canbeih-awn iiithe flowprlwtween 
cycles anil spirals except by their mode of succession. More- 
over, aestivation strictly so called bIiwiH be coneemul only 
with the disposition among themselves of the several members 
of one whorl, or of one complete spiral. So the altcmatiou of 
contiguous whorls, as of the tliree inner with the tliree outer 
(lower-leaves of a LUy or a Tulip (the allemalive n^sttvaUon of 
DcCanilolle) , is a matter of phyllotaxj-, not of lestivation. Tire 
latter is pi-operly concerned only with the relations of each three 
leaves to each other.' 

2^3. The proper sstivations may be classiGed into tliose in 
which tlie pai-ts do not overlap, and those in wliidi they do. Of 
the first, there arc two kinds, the open (out. 
aperta) and tJie valvate, both characterized au(l 
nameil by Brown.* Of the second, there is 
j\ one leading kind, the imbricate (a<lopted by 
/ Brown from Linnteus),with subonlinatt: modi- 
fications.' Accordingly, the aestivation is 
said to be 

*" 2o4. Open or Indeterminatt {teal. tg>erta), 

when the partA do not come into eoulact in the liud, so as to 



' Thp Mine &pp1iet to the two wti uf irpnii Knil of petals In Barbcnr, in 
MtnUpernium, and of the petnli in Pnppy. ftc. (3-W). 

' Liniueus, indeed, iins, " .'EitUtaio valmla. ■! pctnU x? cipmuura ii 
glumic grnminii poniuitur," — the name, but not tlic llilni;: the glumca of 
graeset nre not valvate in tiie botanitnl acngc. Su the ipmi at to its proper I 
one may be aaid to originBte with R. Brown. 

' For a brief ditcuwiun of " ^illration and iti Ti'nninology," »« Amer. 
Jour. Sd. icr. 3, x. 339, 1876. 

As to Tuunn, It ia perhaps more eomct to say of tlie irttivalie* tliat It li ', 
imtirinitirr, nmraltiliet, ralmlar, Ac. (irrf. I'mdri'mlrm, rooiii/wd'™, mlmiri; Ac), I 
liut of tltp leave* or piet^ea, ihni tliey arc iflTlnilr, mni^atr. nt!eii(r, te^ ia I 
BitlTatuin : hut audi precision uf (orni will seldom lie atti-liiliHl lu in lialsn- 
Ical ileKripliona, 

»IH, SIB. ningrMimaim "w™ mxUon nf »i> imon""^! t1' 
eiKlc of flnral Imtm Itopala] valtato tn tlie bud i Uio Inner (| 


cover those within. The most familiar case is that of the petals 
of Mignonette and tlie whole genus Reseda. 

2i)i), Vaivate or Valvular, when the margins meet squarely in the 
bud, without any overlapping, like the valves of a dehiscent ca})- 
sule. Familiar examples ai'e aftbixled by the cal3*x of the Linden 
(Fig. 255) ; also that of the Mallow, Rhamnus, Fuchsia, and the 
whole of the several natural ordere to which these belong. A 
modification of this, cause<l by some induplication or involution 
of the cMlges of the individual leaves, occui*s in most species of 
Clematis : in Clematis Virginiana, they 
meivl}' project within {yalvate-induplt- 
cate) ; in Clematis Viticella, the}' are 
conspicuously involute {valvcUe-tncolute)^ 
or vaivate with margins involute. Some- ^ 257 

times (as in the calyx of ceitain Malvaceie) the joined edges 
project outwaixll}' (or are vaivate with re<luplicate mai^ns) , but 
only slightly so. 

25G. Imbricate or Tmbricattve is the general name for aestiva- 
tion (or vernation) with overlapping. The name is taken from 
the overlapping of tiles or shingles on a i-oof, so as to break 
joints or clover edges. It was first applied, by Linnaeus, to 
leaves or scales on a stem, when thickly set and incumbent in suc- 
cessive ranks or heights, the upi>er partly covered b}' those next 
IkjIow. The involucre of an Aster or of the common Sunflower 
is a t3'pical illustration ; as also the leaves of a CameUia-flower, 
the sepals as well as the petals ; and the sepals or outer leaves 

2SS 2JD 960 

of a Flax or a Geranium-flower afford a simpler but similar 
instance, although, ftx)m the paits Iwing nearly of the same size 
and at the same height, the overlapjnng is lateral instead of 
obviously from below. Fig. 258, 259, and the outer part of 
260, also the inner leaf}' circle of 255, illustrate in diagram this 
true and simple imbricative aestivation of a definite numl>er of 

FKt. 2fS6. ValTatc-iiulupllcate flower-leaves (calyx) of Cleroatis Virginiana, &c, 
7Sl. ValTate-involutc. am in C. Viticella. 

FIG. 2.')R-2G(>. Imbricate lefftivation : 2A8. in two wliorlH of three leaven each (calyx 
aiifl corolla); 260. name of Ave leaves in the onter circle, thorn of the inner circle con- 
Tolate; 250, a nln^le ret of three imbricated leavea (in Uie corolla of Magnolia)^ 
alnuMt oomidetely encircllug each other. 



parte.' It is characteristic of it that some parts (one or more) 
are wholly exterior or covering in the bud, and others (one al 
least) interior or covered, at least the margins. Imbric'ative 
lestivation, it will be seen, naturally attends alternate or spiral 
phyllotaxy (248. and see Fig. 242, 243) ; and if it be main- 
tained that these sets of three, five, &c., in tlie blossom are not 
depressed spirals, but whorls or cycles (as may commonly be 
the cose in the corolla, but hardly in the calyx), it is not less 
true that the jjarts are apt to comport themselves in the exact 
manner of a depressed spiral. The kinds of regular inibricatiou 
of alternate leaves, &c., may be specified by the terms or frac- 
tions expressive of the particular grade of phyllotaxy (i, J, %, 
J, &c.). But some of them have received special names, which 
may be cmployetl, as sultortlinatc to the general denomination of 
imbricate. The most important of these are the 

HquilaTU, where leaves override, the older successively astride 
the next younger. The typical instance is that of ancipital or 
two-ranked (i) conduplicate leaves, successively clasping, at least 
next the base, as in Iris, Fig. 217. In what LinniEus termed 
tguilaHl-triguelroui (well seen in Fig. 240, 241), the leaves are 
three-ranked (Iwiug of the J order), and each imperfectly 
coudu plicate. 

Qiiineuncial KsUvation (as in the outer part of Fig, 2G0) is 
simply the imbricate (estivation of five leaves (%), in which 
necessarily tlie first and second are external, the fourth and fifth 
internal, and the third with one mai^n external, where it over- 
lies the fifth, aud the other internal, where it is covered by 
the first. 

Alternative [estivation, as already stated (252), comes tiota 
verlicillate or cyclic phyllotaxy, and the alternation of successive 
whorls. When two such whorls, say of three leaves each (as in 
Fig. 258), are so condensed or combined as to form apparently 
one set or circle of six members (as in the flower-leaves of most 
Liliaceie), three members alternate with and arc co\ered by the 
other Uiree, and this sort of imbricate restivation is produced. 
Jlore properly, the two series are to be considered separately. 
Where the parts are four (as in Fig. 305), the noftnal irabrica- 
tion is decussate, two exterior and two interior. This is some- 

• All the examples referred (o rrnult (mm sliemaic or tpJnil phyllotaxy, 
the former of higher lerics, the latter uf Ihe i < Fin- 20B. 2SD), and of the ) 
(Fig. 200) order. Instead of aeparating (with DeCandolle an<l olhcn) Iho 
J arrangemmt aa iliSeretit in kind from the Imhricale (under the name of 
qumcuneialKiiirBliDn),«eihould count it ana typical enic. Othcnriee the 
i HrrangF^ent might eqimlly claim a generic dla Unction, alio the ], &c. 



times a clear case of binar}' instead of quaternar}', t. e. to be 
counted as two pairs of opposite leaves ; yet it ma}' be a single 
whorl of four, notwithstanding the imbrication. Or these four 
leaves may even, in some cases, be regarded as a i)ortion of a 
depressed spiral, sa}* of tlie § order with one piece omitted, and 
the others adjusted so as to fill the space. 

257. There are various deviations from normally imbricative 
sestivation, esi)ecially where the members are five, occurring 
some in regular but more in iiTcgular flowers, which need not be 
refeiTed to here. One, for which no specific name is requisite, 
is a case merel}' of excessive overlapping in the regular way ; 
name!}', where each piece completely and conccntricall}' encloses 
the next interior, as shown in Fig. 259, representing three petals 
of Magnolia Umbrella. This the French botanists have called 
convolute aestivation, because the individual leaves are involute 
in a manner approaching the convolute vernation of Linnseus. 
Another is the Vexillar^ as in the Pea tribe (Fig. 30G), where 
members which should be external have somehow dcveloi)ed as 
internal, both in calyx and corolla. A third (which has received 
the usually quite meaningless name of Cochlear^ spoon-like, 
and is also that to which most French botanists singularly re- 
Q^'/tct the name of imbricative) is a state exactly intermediate 
between the quincuncially imbricate and the convolute or 


K }l) 


contorted. In it, one leaf is wholly outside, one wholl}" inside, 
and three with one margin inside and the otlier outside. It occurs 
under two modifications, viz. with the innermost leaf remote from 
the outmost (Fig. 2G1), and with it next to the outermost as in 
Fig. 202. In view of the intermediate character, we had 
applied to this the somewhat awkward name of ConvoluteHmhri- 
caie.^ To bring Fig. 26 1 back to the quincuncially imbricate 

^ It would not be amiss, therefore, to name one of tliese modes, viz. that 
of Fig. 261, Suhimbricatef and the other. Fig. 202, Suhconvdute. George 

FIO. 261. Qnlncancial Imbricate mo(1ifle<l towani cnnvolate by one erlgo of the 
■eooml leaf developing Inside instead of oiitnide of the adjacent edfce of the fourth. 

FIG. 262. Convolute mo<Iifled toward imbricate by one leaf having a margin inride 
instead of outside its neighbor. 

FIG. 283. Convolute, or convolutive, or contorted (twisted) aestivation, in diiM^'am. 
In these three diain'ams, the dark circle above represents the position of the axis, tbo 
ilowen being axillary. 


parts.* It ie characteristic of it that some parts (one or more) 
are wholly exterior or covering in the bud, and olbera (one at 
least) interior or covered, at least the margins. Imbrieativc 
(EStivalion. it will be seen, naturally attends alternate or spiral 
phyllotaxy (24H. and ace Fig. 242. 243) ; and if it be inaiii- 
tuncd that these sets of three, five, &c., in the blossom are not 
depressed spirals, but wliorls or cycles (as may commouly be 
Uie ease in the corolla, but hardly in the calyx), it is not less 
true that the parts are apt to comport themselves in tlie exact 
manner of a depressed spiral. The kinds of regular imbrication 
of alternate leaves. &c., may be spccifle<l by the t«rms or tVac- 
tious expressive of the partjeular grade of pbyllotaxy ( j, 1^. f. 
J. &c.). lint some of them have received special names, which 
may be employed, as subordinate to the general denomination of 
imbricate. The most important of these are the 

ISquilant, where leaves override, the older successively astride 
the next younger. The typical instance is that of aneipikU or 
two-ranked (^) conduplicate leaves, successively clasping, at least 
next the base, as in Iris, Fig. 217. In what Linna'us termed 
equilanl-triquetroiu (well seen lu Fig. 240, 211), tlic leaves are 
three-ranked (being of the J order), and each imperfectly 

Qaincunr.ial {estivation (as in the outer part of Fig. 2G0) is 
simply the imbricate (estivation of five leaves (jf), in which 
Docessarily the first and second arc external, the fourth and fifth 
internal, and the tliird with one margin external, where it over- 
lies the fiflh, and the other internal, where it is covered by 
the first. 

AUrmaiive [estivation, as already stated (252), comes from 
vcrticillate or cyclic phyllotajcy. and the alternation of successive 
whorls. Wlien two such wliorls, say of three leaves each (as in 
Fig. 258), are so condensed or combine) as to form apparently 
one set or circle of six members (as in the flower-leaves of most 
Liliaceffi). tliree members ulternate with and are covered by the 
other three, and this sort of imbricate aestivation is produced. 
More properly, the two series are to be considered separately. 
Where the parts are four (as in Fig. 395), the noAnal imbrica- 
tion is dM-ussule, two exterior and two interior. This is some- 

' All tin- i-xampto rtftrrtd to reiuU from sltcmiip or «plr«l phjllouxr. 
the fomiiT at higher ktip*. ihe Uiipr o( Ihu i I Fi(t. 268, 2fte). uid of the 1 
(Fig. 200) order, Inilend of lepirBllng (with DeCandolli' and otiiere) th« 
J amns^in^nl M iliflcreni in kind from the imbrieaie (under the name of 
qiiinininola1e«tir«tion),vreiIiouMeoan[ it MRlypicnl caie. Other <t lie tbc 
1 smngi'ment might equally claim & gencrio digiincliun, >liu tbc |, &c. 


times a clear ease of binar}' instead of quatemar}', t . e. to be 
counted as two pairs of opposite leaves ; 3'et it may be a single 
whorl of four, notwithstanding the imbrication. Or these four 
leaves may even, in some cases, be regarded as a portion of a 
depressed spiral, say of the } order with one piece omitted, and 
the others adjusted so as to fill the space. 

257. There are various deviations from normally imbricative 
aestivation, especially where the members are five, occurring 
some in regular but more in irregular flowers, which need not be 
referred to here. One, for which no specific name is requisite, 
is a case merely of excessive overlapping in the regular way ; 
namely, where each piece completely and concentrically encloses 
the next interior, as shown in Fig. 259, representing three petals 
of Magnolia Umbrella. This the French botanists have called 
eonroiute aestivation, because the individual leaves are involute 
in a manner approaching the convolute vernation of Linnaeus. 
Another is the Vexillar, as in the Pea tribe (Fig. 30G), where 
members which should be external have somehow developed as 
internal, both in calyx and corolla. A third (which has received 
the usually quite meaningless name of Cochlear^ spoon-like, 
and is also that to which most French botanists singularly re- 
tf^ct the name of imbricative) is a state exactly intermediate 
between the quincuncially imbricate and the convolute or 

contorted. In it. one leaf is wbollv outside, one whollv inside, 
and three with one margin inside and the other outside. It occurs 
under two modifications, viz. with the innennost leaf remote from 
the outmost (Fig. 261). and with it next to the outermost as in 
Fig. 262. In \iew of the intermediate character, we had 
applied to this the somewhat awkwarrl name of Ccnroluie-imiri- 
cote.* To bring Fig. 261 back to the quincuncially imbricate 

1 It would not be amut, therefore, to nmmo one of these mode*, tiz. thmt 
of Fig. 261, .SWmArwritrf^, and the other. Yig. '2ff^, S^Mvmnlute. George 

FIG. Sn. QohMnuKUI imhrir^U iiu>!ified tnwanl cnfirolote by one dU^ ^4 the 
neoMd leaf drrefe>|)iB^ htai.^ liwtead of onts^U of th« ai^^^enit edce of the f^arth. 

FIO. 282. Coarolrite m-idlfiel tnvanl imbricate bj ohm leaf haTiof a marfiii iitf»le 
iMfead of ovtvfrle it« nftifhhor. 

FIG. 90. C-WToliMt. or cr-nTolarlre, or vmMruA ''twisted ■ »»f f raiVm. in <liacraB. 
T« these three dfacraott, the dark circle abore rcpcMcnu the poiltktt of the azb, tha 


parts.' It ia characteristic of it that some [tarts (one or more) 
arc vholly exterior or covering in the bud, and others (one at 
least) interior or covered, at least the margins. Imbrieativo 
(estivation, it will be seen, naturally atleuda alternate or spiral 
phvUotnxy (-248, and bcc Fig. 242, 243) : ami if it be main- 
tained that these sets of three, five, &c., ui the blossom are not 
de|)resscd spirals, but vrhorls or cycles (as may commonly be 
the case in the corolla, but hardly in the calyx), it ia not less 
true that the parts are apt to comport themselves in the exact 
maimer of a depressed spiral. The kinds of regular imbrication 
of alternate leaves, &c., may be specified by the terms or frac- 
tions expressive of the particular grade of phyllotaxy (J, }. %, 
|, &c.). But some of them have received special names, which 
may be employed, as suboi-dinatc to the general denomination of 
imbricate. Tho most important of these are tho 

EquitatU, where leaves override, the older successively astride 
the next younger. The tji^ical instance is that of ancipital or 
two-ranked (J) conduplicate leaves, successively clasping, at least 
next ttic base, as in Iris, Fig. 217. In what LinniEus termed 
egiiitant'triqitetrous (well seen in Fig. 240, 241), the leaves are 
three-ranked (being of the J order), and each imperfectly 

Quincunrial Aestivation (as in the outer part of Fig. 260) is 
simply the imbricate a?stivation of five leaves (J), in which 
necessarily tlie first and second are external, the fourth and fifth 
internal, and the third with one margin external, where it over- 
lies tlie fifth, and the other internal, where it is covered by 
the first. 

AUernaiive Kstivation, as already stated (252), comes from 
verticillate or fyclic phjUotaxy, and the alternation of successive 
whorls. When two such whorls, say of three leaves each (as in 
Fig. 258), arc so c-ondeuscd or combined as to form apparently 
one set or circle of six members (as in the fiower-leaves of most 
Liliaceffi), three members alternate with and are w»ered by the 
other three, and this sort of imbricate sstivation is produced. 
More properly, the two series are to Ite considered separately, 
"Where the parts are four (as in Fig. 395), the normal imbrica- 
tion is decussate, two exterior and two interior. Tliia is some- 

' All tlic exrimpk» rffcrrod to ronilt from altomittenr «piral phyllotBiy, 
the former ot liifclior writs, thr latter of ihv i IFitc. SS8, a(i(1|, and of thi^ f 
(FlK. 280) order. InHle&d of eepurstbg (with DeCanilolle >n<l Dlhcrt) llic 
f HrrmnEcnH-Qt as differt^iit in kind from the Imbricaie (undpr tlie name of 
quincuadalHilratioiil.H'i-ihouldcaunt It aiB tjpiralcBsc. Otherwise the 
) Bmngctncnt might i^iusllf claim a generic dislinvtioD, alu the J, &c. 


times a clear ease of binarj' instead of quaternary, t. e, to be 
counted as two pairs of opposite leaves ; yet it maj^ be a single 
whorl of four, notwithstanding the imbrication. Or these four 
leaves may even, in some cases, be regaixied as a portion of a 
depressed spiral, sa}- of the f order with one piece omitted, and 
the others adjusted so as to fill the space. 

257. There are various deviations from normally imbricative 
aestivation, especially where the members are five, occurring 
some in regular but more in iiTegular flowers, which need not be 
referred to here. One, for which no specific name is requisite, 
is a case merely of excessive overlapping in the regular way ; 
namely, where each piece completely and concentrically encloses 
the next interior, as shown in Fig. 259, representing three petals 
of Magnolia Umbrella. This the French botanists have called 
convolute aestivation, because the individual leaves are involute 
in a manner approaching the convolute vernation of Linnaeus. 
Another is the Vexillar^ as in the Pea tribe (Fig. 30G), where 
members which should be external have somehow developed as 
internal, both in cal^'x and corolla. A third (which has received 
the usually quite meaningless name of Cochlear^ spoon-like, 
and is also that to which most French botanists singularly re- 
let the name of imbricative) is a state exactly intermediate 
between the quincuncially imbricate and the convolute or 

K )l) 

contorted. In it, one leaf is wholly outside, one wholly inside, 
and three with one margin inside and the other outside. It occurs 
under two modifications, viz. with the innermost leaf remote from 
the outmost (Fig. 261), and with it next to the outermost as in 
Fig. 262. In view of the intermediate character, we had 
applied to this the somewhat awkward name of Convohite'tmhri' 
cale.^ To bring Fig. 261 back to the quincuncially imbricate 

1 It would not be tmiM, therefore, to name one of tlicsc modes, viz. that 
of Fig. 261, SvbiwbrieaU, and the other, Fig. 202, Suhconvulute. George 

FIG- 2B1. Qnlnciineial iBMcate moflifled toward con volute by one e<lge of the 
r "'^lopfag faiiMe Imtoad of onMcle of the ailjacent edge of the fourth. 

•fljiite mwlWiiil toward imbricate by one leaf having a margin inMde 

4ata^ or MKVOliitlTe, or contorted (twUted) aestivation, In dlafn^un. 
i«% thtdMk drole above represents the poeition of the axis, tbo 


parts.' It is t-haracteriatic of it tliat some parts (one or more) 
are wholly exterior or covering in the bud. and ottiera (one at 
least) interior or covered, at least tlie margins. Imbiicative 
sestivation, it will be seen, naturally attends alternate or spiral 
pbyllotaxy (248, and see Fig. 242, 243) ; and it it be main- 
tained that these sets of three, five, &c., in the blossom arc not 
depressed spirals, but whorls or eyeles (as may commonly be 
the case in the corolla, but hardly in the calj-s), it is not less 
true that the parts are apt to comport themselves in the exact 
manner of a depressed spiral. The kinds of regular imbrication 
of alternate leaves, &c., may be specified by the terms or frac- 
tions expressive of the particular grade of phyllotaxy (J, J. j, 
|, &c.). But some of them have received special namcH, which 
may be employed, as subordinate to the general denomimttion of 
imbricate. The most important of these are the 

£qmtant, where leaves override, the older successively astride 
the next younger. The typical instance ia that of ancipital or 
two-ranked (i) conduplicate leaves, successively clasping, at least 
next the base, as in Iris, Fig. 217. In what Linnieua termed 
et/uilaat-triquetrottt (well seen in Fig. 240, 241), the leaves are 
tlirec-ranked (being of the J order), and each imperfectly 

Quincuncial aestivation (as in the outer part of Fig. 260) is 
simply the imbricate Kstivation of five leaves (j), in which 
necessarily tlic tlrsl and second are external, the fourth and fifth 
internal, and tlie third with one margin external, where it over- 
lies the finh, and the other internal, where it is covered bj* 
the first. 

Altcrnatiee [estivation, as already stated (252), comes from 
vertieillate or cyclic phyllotaxy, and the alternation of successive 
whorls. When two such whorls, say of three leaves each (ns in 
Fig. 258). are so condensed or combined as to form apparently 
one set or circle of six members (as in the flower-leaves of most 
Liliaceic), three members alternate with and are covered by the 
other three, and this sort of imbricate icstivation is produced. 
More properly, the two series are to be considered separately. 
Where the parts are four (as in Fig. 395), the noftual imbrica- 
tion is decussate, two exterior and two interior. This is some- 

' All llie I'lamplei rofenvd (o roBull from ntltmati! or spiral phyllotaxy, 
thf fonniT of liij^her aimei, ilic Ullcr oC ihc ^ <Fig. 2W. 2G0), tnd of ibe | 
(Kip. 200) order. Instead o( »ep»n»ting (with I>Can<loIle uiil otiien) the 
I arrsn^rocnt Ka different in kind from tbc Imbricate (unilcr the name of 
quincuncial lestiTniioD), we ihoald coont it on a typical case. Othorwiae the 
i arrangi'iuent might equally claim a generic dlBlineiJon, fll»o the \, Ac. 


times a clear case of binarj' instead of quatemar}', t. e. to be 
counted as two pairs of opposite leaves ; yet it may be a single 
whorl of four, notwithstanding the imbrication. Or these four 
leaves may even, in some cases, be regaixled as a portion of a 
depressed spiral, say of the § order with one piece omitted, and 
the others adjusted so as to fill the space. 

257. There are various deviations from normally imbricative 
aestivation, especially where the members are five, occurring 
some in regular but more in iiTcgular flowers, which need not be 
referred to here. One, for which no specific name is requisite, 
is a case merely of excessive overlapping in the regular wa}- ; 
namely, where each piece completely and concentrically encloses 
the next interior, as shown in Fig. 259, representing three petals 
of Magnolia Umbrella. This the French botanists have called 
eonrolute aestivation, because the individual leaves are involute 
in a manner approaching the convolute vernation of Linnaeus. 
Another is the VexiUar^ as in the Pea tribe (Fig. 306), where 
members which should be external have somehow develoixjd as 
internal, both in calyx and corolla. A third (which has received 
the usually quite meaningless name of Cochlear^ spoon-like, 
and is also that to which most French botanists singularly re- 
dact the name of imbricative) is a state exactly intermediate 
between the quincuncially imbricate and the convolute or 

r\r\ n, 


contorted. In it, one leaf is wholly outside, one wholly inside, 
and three with one margin inside and the other outside. It occurs 
under two modifications, viz. with the innermost leaf remote from 
the outmost (Fig. 261), and with it next to the outermost as in 
Fig. 262. In view of the intermediate character, we had 
applied to this the somewhat awkwanl name of Convolute^mhri- 
caU.^ To bring Fig. 261 back to the quincuncially imbricate 

1 It would not be AmiM, therefore, to name one of tliese modes, viz. that 
of Fig. 281, SMmbneaie, and the other. Fig. 202, Suhconvolute. George 

FIO. 261. Q i itiif iwU d imbricate mo<1iflcfI toward cnnvolute by one edge of the 
leeontl leaf dt Wtoytiig tavMe instead of outride of the a<Uacent edfre of the fourth. 

FIO. Ml OMVoiafta ■odlfled toward imbricate by one leaf having a margin inMde 
instead of o«lM* Mi Migkbor. 

^"^. VL, CWifiolnta, or oonTolntire, or contorted (twisted) lestivation, in diagram. 

circle above represents the position of the axis, tho 


parts.' It is cliaratiteriatie of it that some parts (one or more) 
lire wlioUy eictcrior or coveriog in the bud, aiut others (one at 
least) interior or covered, at least the mat^ins. Imbiicative 
a'Stivation, it will be seen, naturallj' attends alternate or spiral 
phjUotaxy {2i>i, and see Fig. 242, 24^) ; and if it be main- 
tained that these sets of three, five, &o., in the blossom are not 
depressed spirals, but H'horls or cjelcs (as may commonly bo 
the ease in the corolla, but liardl.\' in the calyx), it is not less 
true that the parts are apt to comport themselves in the exact 
manner of a depressed spiral. The kinds of regular imbrication 
of alternate leaves. &c., may be specified by the terms or frac- 
tions expreasi^'e of the particular grade of pbyllotaxy ({, J, J, 
J, &e.). But some of them have received 6|iecial names, which 
may be employed, as subordinate to the general denomioation of 
imbricate. The most im^iortant of tbesc are the 

Equitant, where leaves override, the older anccessivclj' astride 
the next j'onngcr. The typical instance is that of ancipitat or 
two-ranked (^) conduplitate leaves, successively clasping, at least 
next the base, as in Iris, Fig. 217. In what Linnieus termed 
tijvilaai-triquetrout (well seen in Fig. 240, 241), the loaves are 
three-ranked (being of the J order), and each imperfectly 

Qaincuacial reativation (as in the outer part of Fig. 2C0) is 
simply the inibricuU; wstivatiun of five leaves (g). in which 
necessarily the flrst and second arc external, the fourth and fifth 
internal, and the third with one mai^u external, where it over- 
lies tlio fifth, and the other internal, where it is covered by 
the first. 

Altemalive testivation, as already stated (252), comes from 
verticillote or cyclic phjUoluxy, and the alternation of successive 
whorls. When two sneh whorls, say of three leaves each (as in 
Fig. 258), are so condensed or combinet) as to form apparently 
one Bet or circle of six members (as in the flower-leaves of most 
Liliaeeic), three members alternate with and are covered by the 
Oilier Ibree, and tliis sort of imbrical« lestivation is produced. 
More properly, the two series are to l»e i^nsidered separately. 
Where the parts are four (as in Fig. a9.'>), the noAnal imbrica- 
tion is decussate, two exterior and two interior. This Is some- 

' All die cxamploa referred to result from nltrrnalc nr vpirtl phjllotucj, 
the former of higher spriii, the laller of the i ( Fig. ifill, 26(1). and of iho 4 

iFlg. am) orrtiT. InsteKil of aepHrating (wilh DeCandolte and otliers) the 
arnui|];eTnent ■■ iliffereiit in kinil from the Imbriealc (under the name ot 
qulncunctMlBitivationl.irethonldeoiuit ilsca typicHleue. OlhiTwi»e the 
i orrangcmcut might i>qu«lly claim • -"lerio distinction, alio llie j, &c. 


times a clear case of blnan' instead of quateman\ t*. e. to be 
counted as two pairs of opposite leaves ; yet it may be a single 
whorl of four, notwithstanding the imbrication. Or these four 
leaves may even, in some cases, be regarded as a ix>rtion of a 
depressed spiral, say of the J order with one piece omitted, and 
the others adjusted so as to fill the space. 

257. There are various deviations from normallv imbricative 
aestivation, especially where the members are five, occurring 
some in regular but more in irregular flowers, which need not be 
referred to here. One, for which no specific name is requisite, 
is a case merely of excessive overlapping in the regular way ; 
namely, where each piece completely and concentrically encloses 
the next interior, as shown in Fig. 259, representing three i>etals 
of Magnolia Umbrella. This the French botanists have called 
convolute aestivation, l»ecause the individual leaves are involute 
in a manner approaching the convolute vernation of Linnaeus. 
Another is the Vexiilar. as in the Pea tribe (Fig. 30G). where 
members which should be external have somehow develo|XHl as 
internal, both in calvx and corolla. A third ( which has received 
the usuall}' quite meaningless name of CovMear^ six>on-like, 
and is also that to which most French botanists singularly re- 
^/ict the name of imbricative) is a state exactly intermediate 
between the quincuncially imbricate and the convolute or 


contorted. In it, one leaf is whollv outside, one whollv inside, 
and three with one margin insi<le and the other outsidi^. It occurs 
under two modifications, viz. with the iniiennost leaf remote from 
the outmost (Fig. 2C1), and with it next to the outermost as in 
Fig. 262. In view of the intermculiate character, we had 
applied to this the somewhat awkwanl name of Countlute-imbri' 
cate.^ To bring Fig. 20 1 back to the quincuncially imbricate 

* It would not be amiM. therefore, to name one (»f tlieso modes, viz, that 
of Fig. 261, Subimbriratf, and the other. Fip. 2»52. Suf-mnnJute. Georpe 

FIO. 28t. Qnlocimdal Imbricate mrxlified towanl c<>nTi»late bj on^ enlcc of the 
•aeond leaf derek^ihiip ln9iik> Inntcail of ouNMe of the a<Uac<>nt eilce of the fourth. 

FIO. SB. Conrolute mrMlifieil tiiwanl imbricate by ouc leaf having a mar^rin in»id« 
I— Tf d of oatvide \t» nei|(htir»r. 

FIO. 9BL ConTolate. or c«^nvolntive. or contorteil (twisted) le^tivation. in dia^rram. 
fa tlMW three diacrama, the dark circle above repre«enta the poeition of the axis, tho 


Ibrm, we have only to reverse a single overlappiug on the telV 
hand side of tlic figure. To restore Fig. 262 to the Lionvulutei 
we have only to reverse a single overlappiug at the lowoi 
hand side. Changes like these, or the reverse, are not rare ia 
several species, i>articularly in the eorolla. The aontial form 
and the deviation often occur in diHeront flowers on the aumv 
individual, thus indicating an easj' passage between the imbricfttB 
testivation in the proper sense and the 

258. Convolute, otherwise called Obtolufe or Contortttt. 
Twisted. Fig. 263, and inner circle of ¥\g. 2G0. Here eat-h leal 
successively overlaps a preosiing and is overlapped liy a folknring 
one, all having a slight and etjual obliquity of position, so that 
all alike have an exterior and an interior (or a covering and j 
covered) matgin, and all appear to be as it were rolled up to* 
„— .^ gcther. This ia strikingly so when the parts am 
^^^\\\ broad and much overlapped, as in Fig. 2G4, 
ifff~ 'J } i"'''"*'''*^ ^'^'^ among the forms of imbricate a;stirft! 
\ l\ v~^ ^ tion, and so does Eichler, [larticularly distin^tsbiiq 
^^?^_X it, however, under the name here preferred. 

w occasional transitions would justil^' such claasifici 

tion. But in most eases it ia so uniform, and in the corolla a 
completely characteristic of whole families (such as Malvai 
Onagniceffi, Aimcynacem, Gentianaeeas Polemoniaceffi, &c.), antl 
is so distinct in its nature, that it may well take rank amou 
the primar}' kinds of testivation. As to its nature, it is evideiii 
that while the imbricate motle (at least the ttrnaiy, quinary;! 
&c.) indicates or imitates spiral ph^Uotaxy (some members b 
ing within or with higher insertion than others), the convoluti 
and the valvate (having all the members of the ^ 
same plane) answer to verticillate phyllotaxy, or to wbod 
instead of depressed spirals.' The name which this mode t 

The tubimhriaib! mode hs) two varietii^B, distinguialicd by EloliI 
BliithcDiliagrsinmc^) u aicmiiiv, when the lower or anlerior (i. «, the . . ^ 
next the gnbiending bract or leaf) are lucceHivel}' exterior (ai In Fig: 98t 
anil daantm, when the coTerinK it from the upper aidi^, ■'. t. from tha dj 
next the axlR. 

' 8U11, M thiMe member* of a quincunx which normally pliould be wbdO 
external do wnnctlmM become internal during their develoiinient in tho bn 
Bimlliir chaneeB may be conccireil In chaD0e a quincuncial into a conTOloi 
diKpoiition : but, to effect tlu», three out of the Ave overlapping! woul4 bsi 
to be re*er»cd, 

TIQ. Kt. ConTolutd (alio ckllryj coutor -»tl4D ol a corolla. 


{estivation ought to bear is not yet well settled, but that of eon- 
volute, here preferred, will probably prevail.' 

259. In recapitulation, these principal forms of the seetivation 
of doral circles may be classified in a synopsis. Thoy are : I, 
those not closed :^ open or indeCermincUs ; II, closed ; and these 
1, with the margins not overlapping ^ vaivate; 2, with margins 
overlapping ; u, one or more with both margins covered = imbri- 
cate i b, all with one mai^pu covered, the other uncovered =: 

2G0. Plicate or Plaited, nhcu applied to the flower-bud as a 
whole, is in a somewhat different category. The term is hera 
used for the plaiting 
of a tutie or cnp, 
composetl of a circle 
of leaves combined ' 
uito one body, 
is well marked in 

the corolla of Convohnilus and of Datura, and 
in most of the order to which these belong. In 
Campanula, these plaits are all outwardly sa- 
lient and straight (Fig. IG'i) ; in the corolla of 
most Gentians, the plaits are internal and straight. 
In Convolvulus and Datura (Fig. 266-268), 
the narrow plaits overlap one another in a con- 
volute way, when they are said to be SuperiohUe. 
In the common Morning Glory and some other "" 

apceies of Ipomcca, these plaits are besides spirally twisted or 

' Sec article entitled " j^ilivation and its Tcrtninnlogj," above rcterred to. 
The earlic'Bl name It Olnvtait, given hy IJnnieiu to the kind ot vernation <n 
whicli IHO leaves (canduplicalc onei in hit dia^am) arc put together ao 
that one half of oath is exterior, tlie other interior. That ia juit the mode 
in queelion reduc-cd to a single pair ot leaves, as il ia in the calyx of a Poppy. 
Mirbel is the only botaniKt who luB applied the term to Kstivation, and to a 
utrcle of more than two leaves, and il has never boen adopted in botanical 
descriptions. Il has the diaadvantaj^ that the prcflx ob to botanical terms 
means ohversely or inversely. Coninried (ronionii). in English Twiaifd, is in 
early and ig the commonest use, and it is sometimes expreeaive. The objection 
to it is, thai contorticjn or twisting of the flower-buil often consptcuoiuly oc- 
ean where there ia no overlapping ot edges (aa in man; species ot Ipomiea) ; 
that really no twisting accompanies the overlapping in a majority ot cases 
of tliis lestivation ; and tliat when there is a twisting it is not rarely in llie 
direction conimiy to the overlapping ; ao that the contortion needs to be 

FIO. 26S. Cmn aecllnn ot the eitronelj pllcala or plaited lube of tha corolla nT 
aCaaipanala In tbabuJ. 286. Samaof a OonTolTDltis<Calystecta),Uie pl^taconvolute 
OI sapervolDtf. 

no. 207. Upper part of nncipnnilal corolla of Dalnraj Of its or 




(ftatarted in the opposite diKctioa ; thftt is, the pisits oreil^ 
to tbt ri^t ami mre iiri&teil to the left.* 

201. J>litetima III <heriapp\mg. &c. This is to be noted In the 
terTum>-, quitian% or other Torms of eptnlly disposed imbricatioQt 
also in cMivolQte aod twistied or oontorted Kstirsttoa. It 
may be either to the right (rfctfrvrar) or to the left (timittrorm). 
Tbe 8])iilicatum of this lenn dependa upoo the assumetl jmaitioii 
t^ the obscner, whether outside or inside. We alwa^-s suppose 
him to Btaixl oal^ide, in fnnit of the object : so when the o(~er- 
]a)i|HUg ia ftom right to left of the observer thus placed, as in 
Fig. 26C. it is sinistrorse ; wlien from his left to right, as in 
Fig. 267, 268, dextronr.* The direction is generally constant, 
but in many cases only pre\-alent, in the same plant or the same 
qjecies, or even the same genua : sometimes it is unifonn or 
nearly so throngboat a whole natural order. 

Mparairi; eipr«»»rd. To deacribe the »*tiTation in inch caws u ^.ifnmia 
contwAi rf (rniin''imini torfi (or ID tiniiUr English words ), when tlie ureriapping 
i* to ihr rigfal ind the twiitiiift (o tlie tefl, ia at U»n awkirard and cutiibratu. 
Vimrvluie it a Siting oamt, ai oivaiioiul carlf application lo Ihl* KstJTatioa 
(a* by Juuiea to (be petal* of HalTarisciu), but without deflniiioo in tliii 
•eiue ; il ha* for tnan; jr*n been tteadilj adapted b; the present writer, 
U vmpUijrci by Kchler in Germanj, and has tvceDtljr been adopted bjr 
G. Iteoalow and othen In Great Drilain. It baa. however, the difadrantage 
of having been lueil lij- Unnem to expten the coiling of nngle leavra, aad 
In a niiniHT not whollj ciHigTuou«, but itill with one edge outaide and Ibe 
other InilJc. 

' In our phnueolDg7, dcxtrorBely convolute and siniitrorKly contorted; 
In lliecurrcDtphraH-iiIogyibove referred to, deicronel^ contorted urlwialed 
anil niniitmrH-ly twialedl 

' 'Die rvaiuiia tor adopting thia view (In oppotiilon to the aulboritf of 
M Mul UcCaniltiUt') an: given in note on p. 61. 





262. IxFLORcscEycE, a term vhich would literally denote the 
time of flowcr-ltearing, was ap[>lied by Linn^us to the mode, that 
is. to tlie tlisi>osilion of blossoms on the axis and as respects 
their arrangement with r^ard to each other. AjrriiOTAxr, a 
name formed on the analogy of i>hylloIaxy. and denoting flower- 
arrangement, is a ttetter term. The subject really belongs to 
ramifieation {HZ, 1-1-lC), and ia also concerned with fohation 
and with iihyllotasy. It is most advantageously treated apart, 
immediately preceding the study of the blossom itself. 

'2(i3. In and near the blossom, 
both asis and foliage ver}- commonly 
undergo modification, either abrupt or 
gradual, giving rise in the former to 
Pedunelti and PediceU, in the tatter to 
Bracti and BraelUU. 

2ti4. A Bract (in Latin Braeita) is a 
leaf t>elonging to or subtending a 
flower-cluster, or subtending a flower, 
and differing from the ordinarj' leaves 
in some respect, usually in shape and 
size, not rarely in texture and color.' 
They are commonly, but not always, 
RHluced or as if depaui>crate leaves, of 
little or no account as foliage, but some- 
times of use for protection, sometimes 
rivalling the highly colored flower-leaves 
for show, more olten insignificant or 
minute and functionless, sometimes obsolete (a8inCrucifer»),or 

1 Brads of (he Bnt order Rre Bomelimes oiled ^ora/ Uarrt ( FiJia Jtonxtia), 
or at leut these are not well diitini^shed from bracts. Bui the term floral 
leave* 1b ilescripliTely more properly find usually applied to leaves below 
the t>Taeta or proper origin of the flower-cluiicrs, yet near them, and un- 
like the proper cauline leaves. It is a vague lenn, and is in some danger of 
I>eing confounded (a« it never ihould be) with another vague term, via. 
flower-leaves, or the leaf-like organs of the flower itself. 

mlp, partly cat awaj bslow to d 

> lbs 


K;tcU tlowi.T is suhtcmliHl by {grows from the axil 
t ill b'iic. 27;-i(H), &i.: A iliistt-r of flo«pi-s is siili- 
i >.^>usllivllOllrt ttiiil ivkiroil lirai't in Fifi. 2ti'J. 'i'f). L'Tl ; 
i.>l\>>K>n>l liRictii. imitating; nhitt' jH-tuls. in Fi^. '2'Jl, 
is llii' iiaiiiv givt-ii to such an i>ii<:losiii>; lirjct, or 
vuvos stiwi-issivtlv t.'ii(.-Iu»iiig a flowLT-cliiater. 

l' ;iiiiiK- K'vi'ii to II fii'cK' or spiral cnllcction "f 
rti i-ilii^U-r, as in Conid (Fig. 2fl4. also in Fi^'. 
«ii'Uii<l a siiiglo llowcr, as in llqMiticn and 
.-.(inl iiiili'iX'iwt'iHi' may Lavf lioili a goncml 
■.niv. Olio I'lT till- gt'iu'nil fl(MVfT-cl lister, otiifi-s 
ii,i; liiisu-r*. 'rill- luinw; of involucre is tlii'ii 
..^lit! .'iiv; tlinl oC 
i,.!»^i U* lln* i>artiiiK socomlarv, or iiltiinflte 

'^ntnMi, itiiiuiiiitivo oriirnot) arc liracts nr 
.liiuulc I'lxler. For ixainiilc. in llu' t-li'inIiT 
.:. -• is a linu't. HuMi'iiilinjr cacli iiKliviilnal 
. tsMUct, or I'rai-t of Miromlarv ni'dcr. Iiorno 
.^-«ui-k ita^-ll". Tlif Fn-iK'Ii naturally Iranslato 
. uw Htii-tiivU (i>l. bnittrolfs) : ill liii^lisli, 
MM..C unt th-ttvF (llminiitivc. 
^a, . ^M.* v»\lv\\ C/irrjT, aro <1iiiiiiiuli\'o or 

■^.,-fcl.>i;i-»t>MiTunilI..Tlth<"rfpml, ITI S|.iil]ie 



chatT-like bracts or r-ractlets on the axis (or receptacle) and 
aiiK'ns lie dc»wer? of a dena€r inflorescence, such as a head of 
Compojiiar iirT.i. Fis. 2^7. ivy) : and the name is ako giren 
to an incer series of ibe 

Gli"mi> of Grasse*. Tbe^c are peculiar ehafly bracts or bract- 
Itft* which chiracterlze the infloresttrncc of Grasses and Sedges- 

i'Oo. PtdiBtle Ls the general name of a flower-«talk. that is, 
of an axis or iicia. which instead of foliage, or at least ordi- 
nal rv fo'.'r»ge. sLj'jx-n* a 
fl<:»wer-cIiislerora sinsle 
flc»wer. In Fig. 270. 
each i^uncle (riaing 
fn:»m the axil *.'f an orii- 
narv leaf. arjKl therefore 
ans-werinc to a branchy 



tiear» a s/jliianr flower. In Fig. 277. the [jedunde bears a series 
of flower?, or a flower-cluster. In this instance, each flower is 
t<fme on a flower-stalk of its own. that is. upon a 

I-^EDKXL. Tl:s is the name given to distinguish 
a partial flower- •talk. or. more strict It. the stalk 
of each in-'iivyiua: flc'wer of an infloreacenc*. i Fig. 
277-2r4. t In less simple flower-clusters, with 
ramif.f-ation of two. three, or more gra/les. general 
r»e:luiicle. f^mia: peiuncles. and piedieels have to 1* 
di^iin:^:!??!.^! : the term pedicel is reserved for the 
ulrlmate raiLif ctkTioa. 

N APE :s :be name criven to a peduncle rising 
frr.'Tii :::e zr^'-'Ai-l. as tl:at of nK»st Primulas, of 
I>>i»-'-a::;«.T.. n-j.a::<::a. and the so-caJled acaukrs- 
«-n: or •:»-il!'-^s V:ol.:t*. ^ 

IJha« h:- ' '*s/-k^-;«rje is a name given to the axis » 
f*f :rif • .rv-^-jfrij'^' : t'^at is. the c<«nt;nuat5on of the 
stt.iD ^T i^'-i-T-r-le thni'Tizh a somewhat el«>ngaled 
fl* •w^-r-flv-vr. a* in a ^y-ike <^«f Bin-h or of Ran- 
la:n. Fis. 1*>. i':-'. \VL*:u t-jis ax:* :- short, as in 
a hf^l /Fiz. 2^.>-2S^ . :t i« u^ualh* «!led the 
Re*: efta'Tle. a "w-^-r I al*-:* use*! for the ai::* or cauline- r? 

r^n <f a fl-'-wer. The (v.Titext shi^uld show when re«-ptacle of 
in3"T>rj^iF-Ei<*-. and when rec^-ptacle of the flower itself, is meant. 
R-'th >¥r]oTiZ to axis or stem. 


Fii^ rr:. a 


wi^k » 



2C,{\. Pmition of Flowen or Cluti'ni. FIowit-IhpI^ &•«•»' 
with Iraf-luidM in r»ri;rin. iM)Htti(iti, mid htnictiin*. ti» tin** t \!* i.' i: 
IruM, tliat tilt* parts of Uitli an' Iravi-s or lioiiin|i*«;ii«^ i.f ;. & . « 
cniwdifl in wIumIs or spiniN ii|)«iii a wliort jNirtinii *»( -?. r:. • 
axis: and a.s Ifaf-lmdH an* i'itlii*r toniiiiial or i I '. 7 ■ 
HO also an* tlowtT-lmdH ; ns a Ifaf-I'titl may jjivi' ri-M- i.i a - :,• •■ 
or a coiniNMiiid <:rowtli, i'. f, umy liRiiich a^aiii niid n;;:iiii. ••: ? *. 
ImiiK'li at all, so llow«T-lH*ariii^ hRiiiclii'H. i»r tin* llMHir-**ar r.j 
rxtn'Miitv of a Htciii or hniiicli, iiiav U-ar n Hiii''!!* fliHAir. *■* % 
iimrr or less <t»MiiMMiiid rliislrr. TliiiH. ill riij. -J't',, nii :i\.. .k^ 
|K-diiii(|4<, or iiaktMl hraiirli, iK'ars at a|N'X a >olit:ii\ i1<>h.7 . .n 
Fi;:. JTT. a |nn1uiicIi> iM-aiN a ItMis** cliiMir of |]iiHi'r«. •a* :i ■ f 
wliirli ^prill^M fn»iii tli«* axil of a small fiRic-t ; in Fij. l'*'>. 
a t<*niiiiial immIiiiicIc U'ars at Humniit a dniM* f1o«ii-r-t in.:, r. 
Flowi-rs an* filln-r Nolitarv or in (•Iiiht«'p*. Wlu-n ^■»li!a^^ . ti.. t 
an* naturally witliont bnicts, )H*tn}; Hufitmdttl iii**t4'ail (a« in 1 :^. 
27(1) liv onlinarv toli:i;;4'. 

2(i7. TIh* i'lr\atit»n fitlicr r)f a solitarv f1i>w<'r or a <'hi*it< r "ii a 
]N*<luncU*, <»r of indi\idiial tltiwcrs of a rlii**t4*r on iMili(i-U. i* ifii!r 
incid«*ntal. TIh* tlowcr^ mav U* stalk l«*!«4. i. i*. «'mi7«*. 

2t»M. The KiDds ofliiflon*M*i*iife wliirh liavi* n^ii-ivi**! di**tiiHfi\r 
names an* various, Imt an* all rrdiiciMi* to twfi ty|ii-«. «h:«}i. 
frriHTally well marktil. may sonH'timrs |ia<«s into rat>li oiIht. .ii>-I 
wliirh an* not ran-ly coniUiintl in tli«- same tfim|Niiind int1*'r*«- 
<i'n<*«'.* The two txiH's dlfl'rr in l>a-is a«* di» axillary i'r*»ni ti r- 
minal )»uds ; in the om* the tlowrr^ an' axillary or lati-rul. iii tfi«* 
other teniiinal in n-^iNTt to the axi** from x\hiih ••a<'h lI<iHtr *'T 
its {Militvl ari'it's. Itnt ina^imiH-li as every ttowi-r, i%liat<'\ir il« 
IMi-^ition. i*4 t4-rniinal to it^ «»wn Malk or axiw. it {•• U-tt«T ti» di«- 
tinirni^h X\u* two t\|N*s in t»tlier t<*nn*i. and to name thi-m ilie 

*.'<>'.*. lDd<*flnit<* and Il(*flniti'. or. in ei|iiivah-nt an«l similur trnii**. 

the Im/ttrrmntnf^ uu*\ I^frrminatt'.' Kaeh may Ik* eitli* r ■•ini|tle 

or (*om|M»nnil. It i<« fp>ni the bimpl** that the di-tlnition« nn* to 
1h' iliaun. In tin- foiiin-r ty)M'. the rhaelii** or main axi<> of the 
ihtlore^^-cnct* i«» iiMt ti-nninatitl l»v a llowrr. l»nt lat«Tal a\i-i. or 
|Niliri-ls. are. In tiir lattfr. Uith the main <»r |irimar\ aii<l the 
latiTal or M-i-i»nil:irv ax<-- or Malk^ are -^o temiinat***!. An in«le- 
tiTniiiiate t1*>\ii'r-( hiNtiT mav ^o on to 4|c\eli>|» intern* "le afti*r 

I Iiitttirt •< • n< f .1* )..!• Niri will iii«io*i-I nil !•> <iiiill.iiiil (iti lt:;ll N^:. 
H'lt I r.iiii I . :v «*' i* ,1 iii'>i!i-. iin* n thiir,: Thf tliiiii;* »i»iim tiii>i« t'Ut iiv 
a]-prii]iruti ly tm < :illi-il arr \\'i\\i r-«-lii«tt p>. fur whii-li, if a t:t rii r»l U* 1itii< al 
ii.tiiir I* n«-««K-«l. th:ii •)( .!'•'/•' 'hi'i, ill Kii^li*h .\f>!K»»iiy, iiii;;i:t-«ti«i l>y (iu:Jau-i, 
i- .!• t*iMHl a« aiiv 

- AN' I tiaiiii-il }•>- r.irlilt-r • niiifhi-nflia|rnitnni4\ 3:i. fullowing UuilUud, L c.) 
thf f.\-*^t»f and ihi* ii'dryitu tyiw. 



liiteTDwIe of axis, and one or more loaves (bracts) at each node, 
Lhi-n a fluwer in tlio usil of ea<.-li braot, tmlil ila strength or 
capability is exhatistcl. Or it mny sto]) short with vert' few 
Hovers ; but the uppermost and youngest one will not rcully tcr* 

jninAU^ tlie rhachis (t. e. come from n termuial bud), tlioiigh it 
ap[>ear to do so. (Fig. 272, 277-279, Ac.) The lower 
ilawer-biiits are evidently the oldest, and acoonlingly the first lo 
expand; and the expansion will proceed regularly fn>m below 
[pward : wherefore this tj-pc of inflorescence has Itccn cqIInI the 
A$etiuiing or Aeropttal; likewise thu Ceniripftal, because, when 
tbe flowers are brought to the same leve] or near it (us in I'ig. 
579, 280) by a lengthening of the lower pcllcfls, with or wltli- 
uut relative shortening of the rhnchis, the evolution 
is Been to proceed from cirenmferenoo to centra. 
Tliera is thus no lack of names ; hut, inasniueh as 
the fiiUowing tj-pe is commonly referreil to under tlie 
general name of O/mote, to this lias recently been 
given tiio counterpart name of Bnlryoie. (271.) 

Ii A determinate flowor-clnster (as seen in its gradual 
lopmcnt which is not rarely presented) has the last internode 

1. m-VO ntacnnu of In 

'I^JM DluBFiiiiu or ilidnlto. ilelemiliiBle, t 

.>ftll«nril»c«i.llnKrwciiiai au.* willUrx 
lt»n laMnl Howern dfrrelaplne, romiltic ■ a-Uowernl 
- ' mlm 3-l1nw«reil, uc > pKlt oT 3-Bniiranil ejtot. 

IftcnulnBU, MmlrliMUl, or lKitt]-nH tn 
itrlfngBl. or ejnumii tnflomi 

flal iow 




of its axis terminatefl bya flower (Fig, 281-284), whKh auswers 
to a Itinninal biid. If niore flowers apjwar, so us lo compusc a 
cluster, tliey spring from the axils, preferaiily fVoiu tbe highest 
axils, and arc later. Tbe order of evolution is shown in the figures 
by the size of the fluwer-biids or degree of expansion of the 
blossoms. Fig. 2S1 best shows why a determinate or definite in- 
florescence is sometimes said to be DtKending ; Fig. 283 shows 
wliy it is called Ctntrifugal, the central flower first cx|ianding ; 
Fig, 284 exhibits the lateral or circumferential paitial clusters 
later than Ibc central blossom, and their lateral flowers later 
than their central. 

271. Varieties of Indetermlttate or Botrfoselnfloresconce. The 
names or most of these have been flxeil from the lime of Lionieus, 
but defined without rorerence to the order of pvohilion of tlie 
flowers. Thej' are the Raceme, Corymb, and Vmliel, with flowers 
raised on pwlicels ; tlie Spike and llend, with sessile flowers ; 
also soma modifieatioBs of tliese, notably the Amtnl and the 
Spnd!x. The raeeine niaj' be taken as the tj-pe. Botrys is 
cquivniciil to racemm, &c. ; and, aa the ty|>c includes diversity 
of forms to which the name racemose would seem inapplieablc, 
the term hutryone (botri/fitrhen of Eicbler) is best chosen as the 
general name of tt, and is a good coimtcrpart to cymose for the 
other typo. 

272. A Raceme (illustrated in Fig. 272, 277, and by diagram 
in Fig, 278) is a simple flower-cluBter, in which tlic flowers, 
on their own lateral or axillurj* pedicels and of somewhat equal 
length, arc arranged along a ix'latively more or less elongated 
rliiichis or axis of iufloresecnce. Tlie conunon BnrlR'rrj', Ciir- 
mnt, Choke-Clierrj- and IJlack Chcrrj', and Lily of the Valley 
are rnmiliar examples. 

273. A Corjmb (Fig. 275, 27!i) is a aliortcr and broader 
botryosocliister, whiehililfere froma racemoonly inlhe relatively 
shorter rliacliis and longer lower iHHiicels ; the cluster tlms be- 
coming flat-topjKjd or convex. The eenlriiwtal character ia thus 
made appni-ent. Tlie gi-eater number of the corymbs of Linnipus 
and succeeding lK>tiinisIs are cjTnes. the central flower first cs- 
[landiiig. And tbe tcnn eorymbote or corymb-like is still much 
usi'd in descriptive Iwtany for a ramification which is mainly of 
the cynioso typo, and where in 8tri<!tness the terra cjiuoee should 
he employed. 

274. An Umbel (Fig. 280). as in Asclepias, &c.. difll'rs ti'om 
a corjTnh only in the extreme ahbreriation of tlie rhacbis or axis 
of inflorescence, and the general equality of the ))ediceb which 
thus all apjKar to originate from the ajiex of the l)edunele, and 



80 resemble the raya of an umbrella ; whence the name, and 
whence also tlie i)e(liccts or i>artial peduncles of an umbel are 
terme<l its Rays. Tlic bracts, brought by the non-development 
of iiitei'nodeB into a depressed spiral or apparent (or sometimes 
real) whorl, become an iaoolucre. (264.) An umbel or any 
similar cluster nlieii sessile (without a common peduncle), and 
the parts <;rowde<l, is sometimes called a Fatcicle (or the i>ediccl8 
said to be fascicled) ; but this term has been differently defined. 
(280.) It is better not to use it for any special kind of infloi'es- 
ccnce, but simply in the sense of a bundle of wUate\er sort 
This will aeconl with the sense in wliith it is appbcd to an 
aggregation of leaves. (248.) 

275. A Head or Capitnlum (Fig. 285) is a globular cluster of 
sessile flowei-s, like those of Hcd Clo^er Button bush and 
Plane-ti-ee. The pedicels need not be abeolutelj wantmg, but 
oulj' verj' short. An umbel 
witli iMXIicols much abbrevi- ' 
ated thus passes into a bead, 
as in Kiyngium, &c. And 
a head with rliachlB elongated 
passes into a sjiike. The 
short rhachis of a head very 
commonly takes the name of 
recepacle. (-260.) The whole 
may be subtended by con- 
spicuous bracts forming an 
involucre (2G4) as in Fig. 
280, or niaj- l>e destitute of 
any, as in Fig. 285. On ac- 
count of the compactness 
and mutual pressure under 
gi'owlh, the bmets among 
the flowers of such heads (normally one subtending each blos- 
som) are apt to bo rudimentarj', reduced to little scales, or 
alK)i1ivc, or completely wanting. In the latttr case, the recep- 
tacle is said to be naked {nude), i.e. naked of bracts: when 
tliej- aix' present, it is paleate or eha^g. A peculiar sort of head, 
not undeserving a six-cial name (though tills is not necessary 
in descriptive Iwtany), is the 

AxTEiooit'M, the so-called Compound Flower of the earlier 
botanists, which gives Uie name to tlie vast onler of Compositffi. 

. Cfplml 


Tbe OMitu: ttxaax " reMrmbling a flower." Althongb it hxs &U 
Uk (iumL-Uin vl a true head, tbe resemblanoe to a flower is 

renuiriutbly Ktriking, tbe involucre imitating a calyx, and the 
Htra{»-Nliuiic<I {liguUue) corollatt of the )H.-vcral flowers imitating 
the i>etahi of a Hiuglv ljlo««um. In Honic (Biicb as Dandelion 
, and theCicLon-, Fig. SrtO ) . all 

Uic flowers of the head l)car 
^ thoKG {H.-taI-likc corollas ; in 
' more (Hueh as Aster, Sun- 
flower, and Coreopsis, Fi^. 
287), only on onter circle of 
flowers docs so ; the reiiinin- 
der, smaller and filling the 
centre (or ditk). may liy the 
casual obsen'er be taken for 
Htamens and piHtils, and flirther the deception. The rhachis 
or re(.f-ptaclc of a head <tf this kind is commonly depres.-M'd, 
iHiiirinf^ tlie flowers on what then becomes the upi)cr surface, 
wliieh udik to ttic imitutiun.* 

SrciiNiL'M. This name, jjivcn to the Fig-fhiit, should be here 
referred to, as it w a sort of inflorescence, of the general iialurc 
of a heati, but with rcce])taele external and flowers enclosed 

I The n-ccjiUL-k' of an AnilmdiuiR hai been tfrmctl CUnaiahlam or Phar- 
anthian ; aiii) Hi involucrv, a PrrijAanmthiuai or Ptridinium. Tlic head lias 
Ukcwluc been nuncd & Cepltakmthium. 

Via. DM. Flowering bniich of Clcborr, wtib two hi 
riU. UT. V«UiwlwGU(>no(*limd«raowanataCor 

f llgulaU flowcn, of 


within. See Fig. 657-659 (683) , where its morpholt^y is ex- 
plained and illustrated. Viewed as an inflorescence, it has also 
been named a llTPAxrtiosiLrM. 

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 j'oung 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 eases in 
which the inflorescence is accompanied 
by the i>eculiar bract or bracts called 
a spathc (Fig. 2C9-271). 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 liracts are said to form "" ^ 

a spathc. and ttiere is no spadix. In Palms, the principal reason 
for naming the inflorescence a spadix is its inclusion in a 
spathc before an the sis. 

FIO. SdS. A(1lcenrFlE.3g7.mon«ilKrK«],w1thcn: 
TtADilllIg on Uie rereptiwle. anJ aabMnilHl by lu brer 
Ukdneulnl ny-flowcr ami put of UHUliar (c, c) : Id d 

ilu- perAct Boirer (a)telt 
iMir(b); alim onallEDlMe 
t bncU or leaTwof tka 

na 2S9. Catliln of WbtU BIrcb. 00. Tonng iplke of Plutkgo major. 



AmsT or CxiKn. This b nenly that kiotl of epOcc wlb 
aealy bncu bone by the Birch (Fif. 289),F^:filar, WIOow.ukL. 
M to one MTt of flowera, hy the Oak, Walnot, uhI Uk-kory, 
which are aocordii^y ctOed mKMbMMNa (ran. Calkms miullf 
CiD oir in one pieoe, ifier tameattg or fruttb^. All tnie r**Hnit 
ue nnisexiul. 

277. Any of th»e fonnsof rimpleiiifloicKencennj-beoora- 
poiindc'tl. RacraM»iiiiiytfaemaelresbeiUspo»ed inneeiBeSi^iiike* 
in spikes (as in Tritkvm), heads 
be aggicgaUd in beads, mnbeb in 
Dmbds, corymlis may be corj-m- 
bosely compound, &c. ; fonniog 
compattad nuxmrt, tpUxt, itmM*, 
and the like, the terminology of 
which is ^nsy. The taoet asnai 
case of truly homomorphous com- 
poonding is that of umbels ; the 
itiflorescrncc of mncb the larger pait of L'mbellifer« being in 
oompouud uubela, as in Fig. 2»y. There is then the general 
umbel, tbe rays of whii-b Ifecome petlnncles to 
the partial umbels, and the rays of the latter 
are peilii'cls. i'mbella and UmMtala ilesig- 
nulo in Latin terminology- the general and its 
{■artinl umbels. UmMUli (eoiucd by the Iat« 
Dr. Darlington) may well replace the latter 
as the English diminutive. But umbels are 
sumetimes ruccmoselj' arranged, as in Aralia 
spinosa. beads may be arranged in spikes, 
and so on. 

278. A Panicle, of tbe simple and normal 
sort (as illustrated in Fig. 291), \a produced 
ivhcu a raceme becomes irregularly com|K(und 
by some (usually the lower) of its pe<lieels 
developing into peduncles carrjing several 
flowers, or more than one, or branching again 
and again in the same order. But in com- 
I>oun<l clusters generally the secondary and 
tertiurj' rum if) cations are apt to differ in tj*pe 
as well as in particular mode, giving rise to 
•n Iiet4'romorpbou8 or mixed ui florescence. (2»8.) 

As I.innu'US (lellneil tln' term, anil as it has generally been em- 
ployed In Iwtanieul descriptions, the panicle is a general term 

rill. ZIW> Coraiiuii 

201. A iluiiili p&oklu. 


for any loose and diversely branched cluster, with pedicellate 
flowers. It is therefoi*e difficult to restrict it in practice to the 
indeterminate type. 

279. Varieties of Determinate or Cjmose InHorescence. The 
plan of this type has Ikjcu sufficiently explained. (270.) Its 
siini)lest condition is that of a solitar}* teniiinal 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, lacing 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 i)e(luncle has 
develoiwd from the axil of each leaf of the uppennost pair, or 
where with alternate leaves there is a single uppermost leaf, and 
then only one such i)eduncle, and thus is produced a three- (or 
two-) flowered cymose cluster. The flower of the primar}' axis 
is marked by its bractless peduncle (therefore a jxHlicel) ; the 
lateral and secondary peduncles are known (commonl}' or nor- 
mally) by their bracts or bract; the portion below the bracts 
is proixT iKHluncle ; that above, of single internode, pedicel. 
Bracts, like other leaves, have potential buds in their axils ; these 
in an inflorescence give the thin.1 order of ramification, each 
branch tip|K»d 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 cymes, by itself or 
as a part of a lai-ger cyme, may be calleil a Cymule, The regular 


cyme usually accompanies opix)site or other grades of verticillate 
leaves, but is nut rare in the alternate an-angement. It is 
readiest undoi-stood in an opi)osite-leaved plant with regular 
opposite ramification, as in an Arenaria. Fig. 292. By its con- 
stitution, a cyme proceeds fmm simple to compound. It mat- 
Fin. '£\ti. DicliotoniouB or liiparouB cymo (cyme blpare of Bravals, Dicbosium of 
Eichler) uf Arenaria Micbauxil. 


tcrs Utile whether its tlovelopmont is progressive, the flowers of 
tliL' ultimu1« rami Heatio lis expanding aft^^r tbeearlier liave matured 
fiiiit, uiul will) aiibtending bracts eonsjiinious ur fuliaccous : or 
wiit^tlicr, as in Eliler and Hydrangea (Fig. 293, and in Fig. 
273). the bractii are minute and caducous or aboitivc, and the 
ntmiflcation 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 c^tmc may be properly said to be comimund 
when tlie primar>' axis in it is a peduncle instead of a pedicel, 

ftnd snpports a cluster (e3Tne or cymule) instead of a solitary 
central Uowcr at the main diriBions.' One form of the regular 
cyme, on acconnt of its compactness, is named the 

GLoMEiifLE. This is merely a cymose inflorescence, of any 
BOrt, which is condensed into the form of a head, or approach- 
ing it. Of I his kind is the so-called head of Cornus florida, and 
of the herbaceous C. Canadensis (Fig. 2M), which shows the 

> The dicliiilonioiu or Iwo-bntnrhed cyme a tlie commonrat, but U some- 
timM nmrkprl by >u|iprcuinn of internoiki; ta. tor example, where the 
br>nthi?a are MppRrcntty fn foun, in hn umbclliform way ; bat thc«e Rrc two 
•eta of two, wlih the internoile between llie p»ir» exireniely ihort ; or where, 
Ma in KMer, ihe braiicliei or rayi arc five, in lhi« i^aic i.*oniiating of the sBnie 
two pairs tLDd a central one, which it a many-floweM contin nation of the 
primary axis. Or fi-rnycd cymea, iie., may lie founder] upon altemaie leaves 
with ihortcned internodea, the rays or peduuulM axiUary to them Itiiu 
brouftht into an apparent whorl. 

Br»Taii dislm^iishcd cymes •■ ntiilii/uirnaii. wilh tlirev or more lateral 
Bxea; liipatwu, with two; ami uBj/mr™*. with only one (cyme muliipnrc, 
bipare, unlpare)' To tlieae KicliliT irivM the an litis mi ve names, sev orally, 
of I'lnnchtisiam, Dirtaiiam, and MoRocliiuium. Only the latter needs illiutra- 
tion ; the other* being a* it were compounds of tiiis. 

FlU. 293. ComiioaBd criDaotBjilnncBai wIlL HOB Daatnl and tnlai|odmB> 

Elunl lluwets. 



cotn]wsitioD best, on close oxamination. A condonscd but less 

capitate cyme, or cluster of cjmes, wa9 calle<l by Keeper and 

DeCniidoUe a Fascicle ; and tiiis terminology hag been much 

adopted. It ia proiwrly enough 

said to bo a fascicle, which, as 

used by Linnntus and others, 

means n liundio, or close collection 

of jjarts, Klictlier leaves, jwdun- 

cles, or flowci-s ; but a fascicle is 

not necessarily a cyme (274) , nor is 

thoi-c nce<l of a special substantive 

name for a comiMict cyme, whicU 

may either be simply so called c 

it may pass into the glomcrule. 

281. Botrjoldal forniH of (^inose 
Tn>«, or False Kocemes, &e. The 
regular cyme seldom continues 
with all its ramifications. 
Fig. 2'J'i, after tlic second forking, 
one of the two bteral pedimcles < 

mostly fails to ap[iear. and in some »* "° 

parts one of tlie bracts also ; and ultimately the lateral poduneld 
present is bractlcss. like the central, therefore equally incapable 
of further ramification, tjcing reduced to a pedicel of a single inter- 
node. This suppression some- 
times l>cgins at the llrst fork- 
ing or at tlie very base : and, 
when followed tlirou<:hout, it 
reduces a l)it>arous ordichoto- 
mous c>ine to one half, and, 
converts this half (when the 
axis straightens) intotlic sem- 
blance of a raceme if the 
flowers arc |>cilicellwl, or of a 
spike when they are sessile. 
Fig. 2ilG is a diagram of such an inflorescence as that of Fig. 292. 
with one lateral branch uniformly suppressed at each division, the 
wanting meml)er8 indicated hy short dotted lines. Cases exem- 
plifying this occur in jwrtions of the in florescence of some of our 

Bnwera; I1i]>isul>(c 

Cnrniu Can»>l«i>ls: Howerlnic *tem Imrloe 

Into • pcluncle, and trmilnaled by > glnmeralB ot rery amsl^ 
[| br a i-rdorol bihI cnrollk-llke iDToIncra ol Ibar liracU. SSS. One 
iin thv gtgnierule, enlarir^ 
uacymaor lyru podlal tMH nesme, 
tb» Mm*, wttb ftlunute leant oi 



smaller H;^-pcriciuns, and notably in H. Sarothra, in nliich the 
Ifavi-B are all reduced to brocte. It u not alwaja easy to show 
why this is not n true raceme. But the other broit ufthc pair, upon 
lliut su|)|)oeitiou. is unaccountably empty : the successive angular 
divcrgeueeofeatli joint of the asia of inflorescence in tlie younger 
part, whieli commonly runs into aeoil, finds explanation in the \'icn- 
tliat each portion is the lateral branch from the axil of the subtend- 
ing leaf: and occasionally the other osil produces a simibr one, 
tints revealing the cymose character. When the bract from the 
axil of which tlie missing branch should come disapiieai-s also, as 
sometimes it does, and uniformly on tlie aame side, a state of 
things like that of the upper part of Fig. 297 occurs. The same 
figure may ser^'o for the arrangemeut corresponding to that of 
Fig. '2<J6. only with alternate leaves. But then, close as the inii- 
talion of a raceme here is, tlie position of each flower in respect 
tx) the bract supplies a criterion. ^\''hile in a true raceme the 
flower stands iu the axil of its bract, here it stands on the opi)o- 
sitc side of the axis, or at least is quit« away from the axil. 

282. Sfmpodial forms. The expknation is tliat tlic axis of 
inflorescence in such eases, continuous as it ap|Kars to be, is 
not a simple one, is uot a monopode, but a i^mpode (IIO, Il(i, 
notes), I. e. consists of a series of seemingly supei'ixised inters 
nodes which belong to successive generations of axes : each axis 
bears a pair of leaves (Fig. 21IG) or a single leaf (Fig. 21i7), 
is continued beyond into a peduncle (or pedici'l in these 
instances), and is terminated by a flower. From the axil 
promptly springs a uew axis or branch, vigorous enough eoou 
to throw the adjacent pedicel and flower to one side : this 
bears its leaf or pair of leaves, and is terminated like its pn'de- 
uessor with a flower ; and so on indefinitely. The fact that the 
alternate leaves or bracts are thrown more or leas strictly to one 
side and tlie flowers to the other. In f^g. 297, shows that these 
leaves do not belong to one and tlie same axis ; for nltematti 
leaves are never one-ranke<l or disjwsed |»rei>onderatiiigly along 
one side of an axis, as in this diagram, and as is seeu in the 
inflorescence of a Houseleek, Ac. 

283. A further difficulty in the moipliologj- of clusters of this 
class comes from the early abortion or complete supjiression of 
bracts. Tliis la not unkuown iu botr^'usc inflJorcscence, occurring 
in the racemes of almost all Cruciferw : it is veiy common in the 
c^tmose of all varieties, and cs|>eclally in the unipnrous ones in 
question, which characterize or a1>ouud in Borraginaccis. Hydro- 
phyllnccie, aud othrrnatural orders. In some genera or species, 
the bi'acts are present, or at least the lower ones ; in otlicrs. 



absent ; in some, either occasionally present or wanting in the 
same sixjcies or individual. It is onl^' by analog}', therefore, 
and b}' 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, these 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 quaUfication that they 
are false racemes or spikes^ or cymose racemes^ and the like ; or 
else, b}' reversing the phrase, with stricter correctness they call 
them racemiform or spiciform cymes ^ &c. 

284. Commonly these false racemes or spikes (or botrj'oidal 
cymes, if we so name them) are circinate or inmlled 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 tliis as scorpioid. 
As the coil is a helix, it has also been named helicoid,^ The 
flowers are then thrown, more or less strictl}*, 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 sj-mpodial and botryoi- 
dal forms of cymose inflorescence become rather difficult ; and 
the sorts which have l)ecn elaborately classified into species 
(and have no little morphological interest) are connected by 
such transitions, and are based on such nice or sometunes theo- 
retical particulai's, 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 cymes is 
that of Eichler in his BUlthendiagramme.^ 

' Scorpiind and Ihliroid have been cJirefully distinguished by later 
morphologists, on account of some difference in the mode of evolution and 
arrangement of the flowers along one side of tlie 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 
a. Lateral axes tliree or more : Pleiochasium, the midiiparous cyme of 

fi. Lateral axes two: Dichasium, the hiparous cyme of Bravais. 
y. I^ateral axis one: MoNOcnASiUM, the uniparous cyme of Bravais. 

The latter, or the corresponding divisions of the preced' *• may be 

divided as follows : 


286, Sundry coniplieations and obscurilies are occasionally 
etKoant«red in antliotaxv or phjllotasy, wliicb cannot lii'iv be 

• Lateral axPB tnnisrene to the rel>tivrlj main axis. 

1. Laurml axes In tuci-eiiivc geoentiona alwity* falling on the Mmc 
ihIp of the rrtalivelf Ruun ax!« : Schsil-uei. [KruwUkc] or Buststx 
(ringlet or curl], the uitiparoai kdicoid cgmt of Bravaig. 

± lAtenl asea (ailing alternately on opposite ei<les of the rclntiecly main 
axit: WicKEJ. or CiitciitxiTS [a carl], the utu'parma iforjiioid cgmt at 

• • Latoiil axei medial (in the lame plane) relative to the main axia. 
& i«ml axn in laccesiire generations alwayi on the bat^k side of the 

Kik from vhich it springs: Fachel, Rbifidich |fan|. 
4 I^Mnl axe* in lucccnive generations always on the upper side at the 
ax^ Itwn whkh it springf ; Sichei., DimpAMtuii [sickle]. 
TW tnbjoJiMd simple diagrams from Gichlcr (Fig. 208-303) illuitrate 
ifcge te«n. The ramiflcalion is given without the bracts, which Ihcoreti- 
^tf «r «ct»dly (Ubtond the axes of each generation. The iludeiit may add 
ihH. 1*4 nt nore readily apprehend the cliaracters. 

BAk-r mMgniics the forms with median (antero-posterior) position of 
axes in Monocotyledons only. 
It is natural to distichous phyl- 
lolasy, and it ac'ords with 
the genera] rule thai, in mono- 
cotyledonnus plants, the first 
leaf of the branch, or Itie com- 
monly solitary bractlet of the 
* pcdmicle, stands over against 
and facing llie bract or leaf 
^"^ from the axil of which said 
posterior and next the parent axis, a* 

^-ij.o->*- CX<^g> 


o ( o) 

Ha. INnemm nt llie Dattryi Tlie Q< 
, XQ. (ironnil |>lu. liiille&tJng iIm an 


explained, except throagh full details : Buch as flowers standing 
by the side of a leaf, or a small leaf by the side of a larger one, 

The transTcrse or oblique position of secondary axes or peduncles, as in 
Eichlcr'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 Khipidium and Drepanium, but on one 
side of this plane or the other. This is most common in Dicotyledons (in 
Drosera, Seduni, Sempervivum, and Ilyoscyamus, in Borraginaceae and 
Ilydrophyllacea;, &c.), and is not rare in Monocotyledons, especially with 
tristichous phyllotaxy, as in Tradescantia. In the Bostryx, Fig. 200, the 
bractlet is anterior or falls on the same side as the bract, or, in other terms, 
the successive l)ract8 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 arc commonly one-ranked. In the Cincinnus or true scorpioid 
cyme (Fig. 208), and equally in the Uhipidium (Fig. 300), the bracts fall 
alternately on opposite or different sides of the sympodial rhachis, because 
the single bract {I/) of each successive secondary axis {a') stands next the 
axis {a) and over against the bract {h) 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 duster, they arc usually sessile or nearly so (or spicate), 
as is well seen in Heliotropes, and in very many Borraginaceous and Hydro- 
phyllaceous species, in Ilouseleek, Tradescantia erecta, &c. Tliis comes 
through antidromyy 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. *J0(] 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- i if i 
cies may as well Ik; reduced t6 two. As these A V/ / "^ \ 
severally include the scorpioid and the helicoid i ^ ^ V/ V 
uniparous cymes of modem anthotaxy, these Jl^ '©) 
terms may he retained to designate them. Or, 
if other terms in use be preferred to scorpioid 
and helicoid^ the form with two-ranked flowers ^ 
may be denoniinate<l Cincinnai, that with single- ans^ 
ranketl Bttstrtfchoidal. 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. iWO 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. 

FIO. 305^. Ground plan (firom Eicbler's Blfitbendlagramine, !. 38) of the scorpioid 
inflorescence of Tra<lescantla erecta, between bract (B) below and axis (a) above: 
I.. II., III., &c., the Buccewlve flowers: p* Im tbe bractlet of the flrst and bract of tbe 
second flower, and no tbe others in snccesnion np to r^ and a small undeveloped one 
beyond. Tbe figure 1 affixed to each flower indicates the first floral leat 


f»r n |)r<li<vl or iNMlnnclr »1n>v«* aiul out nf <'«itifi««'tiiin m.'i. • .■ 
Ifaf wliirli slioulit Mil>t«*ii4l aii'l :i(t*i»iii|»:iiiv it.* 

'2x7, MKed InflorrMTftirt* is tint iiiiiiiinriHiti. TKi^ i. .' . 

p\i'II to rliiMiTsor nilllitlratiuils in whirli tin- l»ii|\|-'* :»:■ ■ 

juiiiiMt. Il4*iii;r liftnidiiori till Ills, tliry an* aliii«*tt ii«i • oo.ii..\ • - . 
iNtiiml. tilt* twii tv|M's )H*ltiii;;ili;; toitiirrri'lit onli-rs of r:irit f. t' 
Hut iiikKt it inav !h* iiirin^lctl caM's of «'«iiii|iar:iti\i !\ • 
iiitlun-scciirr. at IraM in tli«* iH'^iiiiiin:;. S4>iiii' of hIih u t- 
I'lisf the two i\)N-N into oiir. In till* TraM'I (l)i|»**a« u-). :i(i ... , 
fntly siniitli* lii-a<l or short spiki* (tmifs tirnt int«» t\*>^% r ..* ' • 
midiilf. from wliirli tin* llowrrin;j |inMn'*iIs n*^nlarl\ t'» !.♦.« ■ ;- 

llail it U*<^nn at tlu* to)». it woulit nnsw«T to Fi;r. *.'**!. « 

!ilo>soniin;; fnun alMiv«> ilownwanl !»y simple unitli*PM> !..*' -^1 
axes alon;z a niono|MNliaI primary axin, is n siniplr rati ni.:' ":: 
<*vmr. wliiK' it mav also Ik* rall*'«l a rfrtrgrti or tirtrr minute r-r*-'.?** 
Sdmi'tliini; of this si»rl may U* mth in ii'rtain fi|N-i'ioi« i-f C -ir:,. 
paiinhi. with viniatr intlitri'M-rntv. tin* terminal !iloi«-*iini lar! ••!. 
(Ill* olht'r*« tollowin;; irn-^ulaiiy. or partly «townwanl aril |.i:t.v 
upwanl. In ('. rapunt-nl<ii<li>**. wlu-n nitlu'r «li'pau|i4 rat*- :):«•! 
till- inflon-scfmv hiniplr. tli«* r\ohiti<in i<* that of a trm* r:ii^ :ik . 
cxri'pt that a fl<»wrr at Irnirth ttTininatt-s th«* axis an«l ili\fl"|'« 
caiiirr than tlir np|N'r half f»t' the nui-nir. In Liatn<« •>|k<:i!:i 
an<i its nt'ar n-latiM-s. tin- hfads, on th«* xir^ati* tri-n«ial riXi*. 
come into lliiwir in an alnioM n-^ular (lrs<'«'n«IiM*^ onhr. "r ar« 
rvrirstltf %fii*ntt\ \\' in Fit;, -■'s] tlu* lowrr jM-iliii-N wwv pr»'i«'rij»'»l 
to till' h\i 1 of thf npiN-r. a sinipit* ror\ml>itorin rwiir w«ii:!>l U- 
M-4-n. with '^inipli* rrntritn;j::il cvolntion, that is. r«-jiil:irU t'r'*m 
till' i-cntn- tiithi* margin : lhi^i<« the ronntrrjiart of tht- rhipi<l:ti!ii 
or r:in--»li:iiM -1 < \ ni«*. of Fi::. :inn. In whirh th«* r\oIntion of iIm* 
|ilii^^i>ni-^ \^ i\^ ri"jnl:iil\ rtntrijN-tal. Tlu* fxplanati«>n of \hv 
parM<lii\ i^ n"t far to >r«k. 

' 'I 111 |i-t«-*: 'I .f ;i p. ■!■■ I I Ht Thi- •i-li- nf .i !ir:|i ! in f.i!"i' fU'-t Hh • !• • 1 
jil.iint 'I Ml ;!:• ! ■•■ J iiiL' ii-rr. l! iii:i\ <H-i !ir Ml trui- rail I !»•»*« inrt'-ri ••« *!■ »■ 

l>y !li< ri!:i-'.-:i •■! >.--:.• ii>!.irv r:ii • iiii-* tlnun tn .111 unit* I nf :« > 

It iw. r«. f : t:.-^ • :•«■ 1i t'l- '■' i- ! ij» ill iii.iii\ -jn % ii ■ nf iK •:i.>i-ti'int '. am I 
tliu* «. . ti.:'..*\ l.iT. :■■ i*. 'ir !■» a •inu'I' M-«»ir fU tin n^'li! "f li f* "f :! 
'Mm- I ■ I'.i •■ I :• • ■■! .1 ji' !' • I !•• ttii- ;i\i» fi-r :i i«in«i'lt r.i!»|t lit ij? ! at««%t :lif 
• nil!, ii I M_' \t \' '. '.'.\ .1 » !i I !•■ ir.f!"rt •• I !'•■■■. nr iilnxr tl'i- |.i*t li af ip a 

^\ inl"' 1: il "• . ■ f s. J.iujjii r . i« rniniiiiill Ni I»k» » >• l-rai t» 

or I'lV'- i:.i\ '- I -r -i 1" ■ ■ 1 il.-Vi'- . a-In.if' I-i -i iniH..l .il •Ii«miT«. »!h tfn-r 
)m 't'.ii. Ii « ir '.' if> !!■%*•:!•••'l'i •. Tli:* >ii:inM--t '•• j':-!-" » l»T 
Si ' I'lj.. r :•■?■. ■-• Ir. -J .. • r ■■iiMirr' H' • in i .r 1 ni iKitiira. .Vir i».i. 
n.- -t -^ I . .i * ■! s- ! ii-'iMi. \. r, :iii.l i« !l.t' I \]>! in.i!!i»n nf itn-ir •■»^aiI«J 
■■'• li.i\i*. w f.i ri a 1 ir.'i- li:if riiiily t.« Ii!i;:in;: ItiWiT il-iiin \\x* a 
^\u.\\\ 1. if l.v :!.. •'. !• ..f !t. N. \V\.|1. r sn M-it Ail. ii Ox*. ic, Stiiiiiinr 
in J I ltra«. & l-:, an.l Kiiiilir. Uiuihiud- i r/.*. 


2S8. CompoutnJ mixcil infloreBcenoe is vorj" vflriotis ninl com- 
mon ; litiL Uif c-oitiliiniitions have hardly callwl Tor sjuHial ti-mis. 
I>cing usually di§poso<l of by a 60)>arato inerition uf the g;cncral 
and of the partial antliotnxy. nr Uint of tin- main axis of iiiflo- 
resci-iiM' and lliat of its ensuini; ramification.' Id C'omposiUi;. 
ffir instance, the llowcre are always in true heads, of eenlripeliil 
pvoliiriun. The beads tcnnina<<.- iii}iin sixms as well as latcinl 
(>nin<;lii.-s, bo that lliey are oeiitrifiignlly or cjnnosely disposed. 
Tlip reverse occurs in all Labiate and most ScropUulariaceee, 
«berc the floivere. whcu elii8t<?re(l, are in cjnnt'B. hut tUcse cjinea 
are trvm axils, and develop in centri|KtAl order. Jt is tliis 
arrangement which mainly characterizes ihe 

'I'limsi's. A compound intliireseenee of more or less elongated 
shape, with the primary ramilicnlion centripetal or bolrjosc, the 
secondary or the ultimate jcentrifugal or cj-moae. To the dcfliii- 
Uon i» geiicndly added. tJiat the middle primary brunclies arc 
longer than tlic upper and lower, rendering the whole cluster 
naiTower at top and Imilom, and Bometimes tlial it ia compact; 
hut tJiL'sc particulars belong (mly to t,vpical examiiles, such as 
the inltoresceDceof Lilac and Ilorscchestnut. In Itiefoi'mer. Ihe 
thjTsiUf U usually eom])ound. A loose thjTsus is a 

MlXEP I'asivls. It is seldom that a repeatedly branching 
inflorescence of the paniculate imxle is of one type in all its 
Miceessive ramifications. Either the primarily centripetal will 
beeonie ceiitril^igal in the ultimate divisions, or the primaiily 
ccntrifhgal will by suppression soon run int*) false tiotiyose 
forms, into aptiarent racemose or ei>ieate subdivisions. So that 
the name P'lnietr in lcrmin<)lo}i;y is generally applic<l to all such 
mised eomitouud inflorescence, as well as to the homogeneously 
botrjose. (278.) 

Vkuticillasteh is a name given to a pair of oiijmsite and 
sessile or somewhat sessile cymes of a Uijrsiis or Uiyrsiform 
inflorescence, which, when full, seem to make a kind of verticil or 
whorl around the stem, us in very many Labiatie. The name 
was originally given to each one of Uie pair of cjTnoa ; but it is 
belter and more commonly used to denote the whole glomcrulc 
or false whorl [irodiieed by tlie seeming confluence of the two 
clusters into one which surrounds the stem. 

1 Gunku<1 {in hi* ineniiiir on Tnflores<?enGF. puMUIicI in Bull. Soc. Ilctt. 
Fwncp, I*.) pm|Mi»ci to ilc«igniile lu f'ym^Balryrt Ihv mixi'O liiflnn'Eceiire 
oompoKd of cymes dpvclu[«tl in botr^oae order, i.r. tlic thynus; and 
ftj*rj.('ji»fi, till* nrverw M»e nt rncftiif*. &c.. cymosety aggT^gatcd. For 
the fomwr. thf old name tliynua tvrtm approvriately •nd well. 



2H0. The Relations of Brad, Brartlol, and Flower ithouUI here 
be noticed, although the subject in part belonga rather to the 
section on Floral Symnietiy. (315.) 

290. Anterior and Po»t«rlor, otherwise called Inftrior and 
Siiptrior, anil therefore Lower and Upper,^ arc prinmrj' rebtiona 
of position or an axillary flower with reBjwct to aulitending bract 
and the axis to which tlie bract pertains. Tbe flower is placed 
between Uie two. The portion of the flower which faeea the 
subtending bract is the anterior, likewise called inftrior or ioieer. 
The opposite portion which faces the axis of 
(" inflorescence is tJiP poilerior, or superior, or 

upper. The right and left sides are laleral. 
(Fig. 304, 306.) These relations do not 
|l\^api>ear in a solitary flower teiTninating a 
]] I simple stem ; but when such an axis produces 
axillar}' branches witii n terminal flower, the 
relation of this flower to the preceding axis 
and its leaf ia manifest, Just as in indetermi- 
nate inflorescence. 
2iH. Median and Traosverse. The jioaition of parts which lie 
in ante ro- posterior line, or between bract and axis, is median. 
Thus, in Fig. 304 and 305, the parts are all in the nicdian plane : 
iu Fig. 306, tbe bractlets, b", b", are taieral or eoUatend, or (being 
in the opposite plane) Iraim-ene. 

S92. FoslUon of Bractlets. Tbe rule has alreatiy been laid 
down (285) that the first leaf of au ullemate- leaved secondarj' 
axis is in Monoctyledona usually median and [xiBterior, thai is, 
farthest away from the subtending leaf (as in Fig. 304, 305) ; 
in Dicotykiions, lateral or traiiaverse. When these secondary 
axes are one-flowered peduncles or jx-dicels, the leaf or leaves 
(if any) they bear arc braellel*.' Commonly there is only this 

I Not (wilh propriety, ullliough the term* hare heea to u*ed| exttrior or 
outer for the anterior, and inltrior or I'nntr for tlie |)o>I«rior po»hioii. These 
lermi should be tetenvA for the relilire poshion oa the axis of •ut'ceuive 
circles or psrta of cirtUi, apirali, SiC. Covering or overlappiug p»rtt are 
exterior or outer in respect to those oTerlapped. 

* Latin Bmrleolit: not that they arc imatl bracts, but bracts of an ulll- 
mate aiU. In axillary influmcrncc, the distinction between bracilel and 
hrairl is obriout: in case of ■ salilHry terminal flower, [here is no iiround of 
ditterence: in terminal or cymoae inflorescence, the differences nrbiirnry; 
but we may mtrict the term bnicllet to the lut bract or pair, 

German bolanials mostly distin^iih between bracts, as a leaf subtending 
a flower or cluster, and bractlels, by terming tbe [urmer a Ikckblna. and Uie 

FIG. a 




id its rslallon to axia 


posterior one to a simple axis in Monocot3'ledons, and two 
transveree ones in Dicotyledons, i. e, one to the right and the 
other to the left of the subtending bract, Fig. 30G, b' b'. When 
the latter form a pair, the}' are perhaps always trul}' transverse ; 
when alternate, the}' stand more or less on the opposite sides and 
transverse. When more than one in Monocotyledons, the}' may 
become either median or transverse, or even intermediate. Tlic 
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 
inteiTals ; ^ 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 
fiower-leaf is at J divergence from one (the up[)ermost) of them 
when the circle is of three, or at f when of five members, or near 
it ; but with many exceptions.* 

A tabular view of the kinds of inflorescence and their termi- 
nology, ser\'ing as a key, may aid the student.* 

latter, being the leaves which the new axis first bears, VorblStter, which is 
also the name they apply to primordial leaves in germination. 

* In accordance with Hofmeister's law ; but (as Eichler remarks) not to 
be explained on his mechanical principle of production in this place becatae 
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. 

» INFLORESCENCE is cither Pure, all of one type, or Mixed, of the two tj-pes 
combined. The Typos are : 
I. Main axis not arrested and terminated by a flower. Indeterminatt^ Indefi- 
nite^ Acmpetal or Ascendinf/^ Centripetal, or 

II. Main and lateral axes arrested and terminated by a flower. DettrminaU^ 
Definite, DeKending, Centrifugal, or Ctmosk. 


162 ANTeOTAxy, or inflorescence. 


1. SiagiU, with Ulenl uea onbraached and tfnniiiBlnl by a Bingti^ flower, mil 

Flowcn on pedlrali, 

Of tomewhit equal lenitthon icnmpiiratlvelyelDngalod axii, BaCKHe. 

Tile lowsr onea longer Ihan the uppiT, and miiii axia abort, . Cortub, 

01 nuarlr equal length on an undeveloped main ixii, . . L'mdei. 

Flowvn >ct>Uc on a very short maiu aits, IIkad. 

Flowcn KM-ile on a eompnrativety elongated main axil, . . . Si-ike. 

A icaly-bracledtpilie bin Ah KMT or 


2. Compoaiui, vrilh lateral axes branched OBce or more, bciring clusleri inilcad of 

Irregolarl]- rs«mo>ely or corymbowlr compound, ■ ■ Pamclb. 
Homogcnrously and rrgtilarly compound, a« 

Raceinu in a nc«mr. Cohpoukd Raceme. 

Corymbs corymbose, Comiiiunu Cobtkb. 

Umbuls in an umbel, CuxitniMi Uhbei. 

Spikes B[Hcate, Cokfouhd Sfike. 

Homogeneously compound, the lecotidaiy ramificatinn 

unliki^ Ibe primary, as Ucadt ractnaie, UmbiU qnktd, 

Sjiiku paiUded, ttc 

L Smptt, with lerminal axis ot each generation one-flowered. 
itimnpailiat, tlie axis of each generation eridently re- 

BolTudintobrinclies, Titrs CVHE. 

Thew more Ihoa two, PUiDckadum or Mui,TiPAKiHia CrNE. 

Tliese only two, . . . Dlchadam, DichoUmoui or BirAiiuUB Cvme. 
Bgrnpodiot, the appan-nliy simple axis continued by 

if HUM*™;™, fall fiortmc or Sptki, Boiryott or UniPABOfs C»ii«. 

Flowers one-ranked on one side of rliachia, HbLKshu Usipabihts OyxI. 

Flowers two-isnked on one side o( rhacbis, Scofu-toiD UMirAROis Ctkb. 
S. Compomirf, with Iirmioai axes lor one or more earlier 
Keneratiuna beating ■ cvma inslead o( a single 
fluwcr, Various sorts of Comfouad Cyme. 


1. Aaom«hm Mmptt, with unbranched one-flowered lateral 

axes (287). such BS . . . Partly bevebsei) Sfikes OB Baceheh. 
S. Cvnpoitml, at variooa contbinatlona, ot which there are 
namci for the subioined : — 
Primary infloretcenoa botryoae, with axis elongating; 

aecondary cymoM, THYBBua. 

Pair of turh opposite rj'mea seemingly coaHuen I round 

the main axis, VeBTicn.UBTEE. 

Panicle with some of the ramilication* cymoae, . . . MiXEu PAntCLs. 




Section I. Its Nature, Parts, and Metamorpht. 

293. Flower-buds are homologous with (morphologically an- 
swering to) leaf-buds, and the}* oecupy the same positions. (266.) 
A Flower is a simple axis or a terminal ix)rtion of one, in a 
phsenogamous 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 
alwa^'s easy to determine exactly where the flower begins, llie 
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 ix^dicel 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 
cither obvious leaves or suflflcientl}' 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 show}' parts, though not themselves directly 
subservient to reproduction, have alwa^'s been accounted as 
parts of the flower.* Between the lowest or outermost of these 
and the bractlets and bracts there are various and sometimes 
complete gradations. The axis itself occasionall}' 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 diflTerent 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, b}' taking 
some particular specimen or class of flowers as a representative 

^ Indeed, the colored leayes, or envelopes in whateyer form, essentially 
were the flower in most of the ante-Linnaean definitions (tha*' ^^ 

excepted), as tbej are still mainly so in popular apprehenskm 



i both complete k 

or pattern, and especially Bomc one which is 
morphologically Biiuple. 

2'Jo. Such a flower consists of two kinds of organs, viz, tlie 
Protecting Organs, leaves of the blossora, or floral envelopes, 
wliieh, when of two seta, are Calyk and Corolla; and the 
Essential Reproiluctlvc Organs, which co-oi>erat« in the production 
of seed, the Stamens and Pistils, 

296. Floral EDvelopes, Periantli, or Perigone, the floral leaves 
or coverings. The former is a proper English designation of 
these (mils, taken collectively. Bnt in descripti\-c botany, 
where a single word b preferable, sometimes the name periaiUh 
(Lat. perianthium) , sometimes lliat of /wnyone (or pen'ifoutam) , 
is used. PfriarUhmm,' a. Linniean term, has lieen objecteti to. 
because it etj-mologically denotes something around the flower ; 
but it seems not inappropriate for the envelopes which surround 
,„ the essential partof the flower. Perigontum, 

a. later term, has the advantage of meaning 
something around the repnxluctivo organs, 
which is precisely what it is. Neither name 
is much used, except where tlie ])eriauth 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 ap])carance of one 
and neeiling descrip- 
tive treatment as sudi, 
ns in tlie petaloideous 
Jlonocotyledons. It is 
also used wliere the 
„ as sa nti m morphology is ambigu- 

ous. Generally, the floral envelopes arc treated disliuctivcly as 
calyx and corolla, one or the other of which (mostly the corolla) 
may lie wanting. 

■2'il. The Cnljs is the outer set of floral enx-elopes. That is 
its only deflnition. Commonly it is moi-e 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 

' Linnaiia (and al«iut the »ame tiine Ludwig) uspil It in the tentc of a 
proper cilyx. yel with Biinio Tagni-nen, Mirbol and Ilrown catiihlifihcil it 
in the wnw ol the collrPti*c Bonl poTcring. DeCindollF n:vi»cd Elirh«n'i 

FIO. MI. Tbecomplctcllovat ofaCruauU. SOB. Ufcram c 
Ib(i bwl, ihcnrlni Uia nlatliB iwdlkm of li> pnrli Thn Svn plwM nt i 
cinja uv laeUan of tlie «pBl*-. tlic oul. of Ui* petKlnj tha Uilnl, of U 
Uimucb tbrlt lUilbcn: III* luBertaad, at Uw lite giliiit*. 

rii). Xin A Hftal; ]li>, * pctali III. ■ ■tuucni; miJ 3U. a pUtll ttata 
r«|>msi>ti)i[ 111 PlK. ytt. 



is as highl}' colored. A name being wanted for the individual 
leaves wliich make up the calj'x, analogous to that for corolla- 
leaves, DeCandoUe adopted Necker's coinage of the word sepal. 
Calyx-leaves are Sepals. 

298. The Corolla is the inner set of floral envelopes, usually (but 
not always) of delicate texture and other than green color, form- 
ing therefore the most showy part of the blossom. Its several 
leaves are the I^tals.* 

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, ser^'e for allurement of insects to the blossom, to 
mutual advantage. (504.) This Aimishes 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 indirectl}' subser\'ient to reproduction. 

300. The essential organs, being commonly plural 
in numl)er, sometimes need a collective name. Where- 
fore, the aggregate stamens of a flower have been 
called the Andkcecium ; the pistils, tlie Gyncecium.* 

301. The Stamens* are the male or fertiUzing 
organs of a flower. A complete stamen (Fig. 311, 
313) consists of Filament (/), the stalk orsupi)ort, 
and Anther (a), a double sac or body of two cells, 
side by side, filled with a i)owder3' substance. Pollen, which is 
at length discharged, usually through a slit or cleft of each cell. 

wcll-fomied name of peritjonium, and in the sense here given. But later (in 
the Orpanographie) he proposed to restrict it to cases in which the part is 
of ambiguous nature, as in Monocotyledons. The earlier definition is no 
doubt the proper one ; but the occasions for using the term in descriptive 
l)otany arc mainly where the nature may seem to be ambiguous or con- 
fused, or where, from the union or close similarity of outer and inner circles, 
it i« most convenient to treat the parts as forming one organ. 

» Fabius Columna, at the close of the sixteenth century, appears to have 
introduced tluH term, or, as Toumefort declares, " primus omnium quod 
sciam Petali vocem proprie usurpavit, ut folia florum a foliis proprie dictis 

2 Tlie male household and the female household respectively, terms in- 
troduced by Ktrper (Linnaja, i. 437), in the form of andraceum Andgynaceum ; 
but the diphthong in the latter should also be a. The orthography 
andrariiim and tjifunrvim (early adopted by Bentham, in Labiatarum Gen. et 
Spec ) is conformable to the Ldnnaean Monacia, Diacia, Ac. 

« The name (from the Greek and Latin name of the warp of the ancient 
upright loom, and thence used in the sense of threads) was applied, down 
to Toumefort and later, to the filaments; and the anthers were termed 

FIG. 313. Stamen, componed of/, filament, and a, anther, with cells opening 
laterally and discharging pollen. 


S02. The PtstilR, one or more lo the flower, are the femat? 
seed- bcuri tig orgaus.' A coiuplete pistil is distingtiisbcd into 
tlirue parts : the Ovary (Lat, 
Ovarium, Fig. 314,0, shown in verti- 
cal section, and Fig. 315 bj' LinuEeus 
named Germeii), the hollow portion 
at the hose which contains the 
Ovules, or bodies destined to be- 
come seeds ; the yrn.E (i), or colum- 
nar prolongation of the apex of the 
"' ovary; and tlie STioMA*(r), a portion 

of the surface of the style denuded of epidermis, 
sonictimea a mere i>oint or a small knob at the 
aiicx of the style, but often forming a single or 
double liue runniug down a part of \l& inner face, 
and assuming a great diversity of appcarauce iu 
difftrtnt phtnts. The ovarj- and tlie stigma are the 
essential parts. The style (its also the filament of a 
stamen) may be altogether wanting. 

apira. It csmG In lime to be used oi now for llic^ whole organ ; but Luil- 
wig {Inil. Reg- Veg.], In 1T42, apparcnltj' first ao defined it, and int rod need 
the term Anlher for tile Apex of Kay, or Tlie<ii o[ Grew. 

I Following Ijnnatui, this tertn is hcK freely used in the plural, and for 
e&ch acti4al separate member of the gyiwEcium, caeh organ which hat an 
ovary. sligniB, and commonly a ityle. Toumefort, wlio appears lo have 
Introduced the word, employed ll in the aente of gyniEcium. Many authors 
deflne it thus, and then practically eliminate from botany this, one of the 
oldest of il« terms, and one by no means superfluous. The typical pisliUaiu 
of Toumefort is that of the Crown Imperial (Inst. i. 90, & lab. 1) and (be 
name is from tiie likeness to a pestle in a mortar. Aa it soon became im- 
pnuihle to apply tlie same name to the piatil uf a Frilillaria or of a Plum, 
the clualer of sui-h organs in Callha, and the capitalc cluster and receptadc 
of sutli organs in a lianunculua or Anemone, Unnteus, and Ludnig before 
him, look tlie idea of Toumefort's name, and used it accordingly. 

" PittiUutii est pars interior ct media floris, qu» ex ovario et stylo com- 
ponitur. . . . Oi-onnm est pan [ualilU inferior, qun luluri fnictus delinea- 
tlonent slslit, . . . Sryfus est pnr« plslilll ex oTario centro produi^la. . . . 
Bummitas atyli rel ejus partlum Sllyma dlcitur," Ludwig. Inat Reg. Vcg. 
41-43, 1742. Without mentioning the plural, Ibc pistil i* thus defined in a 
way which necessitates its use. Linnnus (in Phil. Bot] first defines Stamen 
and Pisllllum In the singular number, cnumeratinil the three parts of tlie 
latter, and afterwards (p. 57) declares that "Fistllbt difFerunt quoad 

Fia SI*. Vsrtlcal KvHun of a phiMl, showing tho Interior of its oTsry, n. to ono 
tdilu of whicli are attaclMid numeroDS otdIm, iI: aboTS is Uie stfle, b, tiiiped bj tho 

Flu. aiS. A (Jstll of Cnuwula, like Ihst nf Pl(. .117, Init man rasgnlflnl. a»l iMit 
acm^iliroHgh Uie o»atT, toshnw IWccll. ar'l the ovules It runtalns; »1« riii11p.l<i|*n 
tietiiw at till Bill lira. At Ilia ■DiiiniU uf tlin ilrls In ■»■• n jtnniDWhat pii|>tll<« porlluu. 
•lalttoteorpi.Mcnnls, citemDnga llttlo wsj-dowii Lhi- luiiet fui' ; ililslt tUostiguia. 


303. The Tonu, or Receptacle of the flower, also named 
Thalamus,' is tbc axis which bears all the other parts, that 
upon which they are all < • 
(mediatelj' or immediately) ^ 
inserted. These are all ho- \\\ 
mologous with leaves. This 
is extremity of stem, or 
floral axis, out of which the 
oi^ans described grow, 
succession, like loaves on 
the stem ; the calyx from 
the very base, the petals a* 

next witliin or above the calj's, then the stamens, finally the 
pistils, wliich, whether several or only one, terminate or seem to 
terminate the axis. (Fig. 31C.) 

304. Metamorphosis. If flower-buds are homolc^ous with 
leaf-buils, and the parts of the flower therefore answer to leaves 
modifieil to special functions (293), then the kind of flower here 
employed in explaining and naming these ports is a proper 
pattern blossom. For the organs are all separate pieces, 
arranged on the receptacle as leaves are on the stem, the outer- 
most manifestly leaf-like, the next equally so in shape, though 
not in color, the stamens indeed have no such outward resem- 
blance, but the ripe pistils open down the inner angle and 
flatten out into a leaf-like form. The adopted theory supposes 
that stamens and pistils, as well as sepals and petab, archomolo- 

numcniin," etc., and ao elsewhere, besides founding his orders on the Dum. 
bcT of piBtils. Among even French authors, Mirbel (1B15) writes, "Le 
aoml>rc Jes piatila n'eal iws le mEme dans touics Ice eip^cea," &c. Moqain- 
Tanclnn freely refara to piatils in the plural, and Aug. Si. Hllaire takes 
wliuUy (be vieH here adopted, distinKuishing the (olituT pistil into simple 
and compound. lleCandolle, in Tii^orie Elementaire, third edition, writes. 
" Choquc carpet est un pcllt tout, nn pistil entier, compost d'un ovairc, d'un 
style, ct d'un stigmatG." Of English authors, no other need be cited than 
KoberC Brown. Tiie terms in question, then, are ; — 

(i^aciiim, the female system of n flower, taken at a whole. 

PiMil, each separate member of the gyD<Edum ; this cither simple or 

Otury, the ovuliferouB portion of a pistil. Substituting a part for the 
whole, this term ia often used when the whole pistil Is meant. 

Ctiqitl, or Carjiid, or Carpo/ihgll, each pisllMeaf ; whether distinct as in 
simple or apocarpous pistils, or in combination of two or more to form acom- 
pounil or syncarpous piatil. 

' By Tnumefort, an<l adopted by Ludwig, Rreeplaeiiliini Jlorit, Linnsus. 
Thona, Saliabury. Tarag (the proper form), DeCandoUe. 

Scdnm tematnin. two of sacb sort, 
c, itunnij c/, pliIlL 


gotis with leaves ; that the sepala are comparativclj' little, the 
petals more, anil the reproductive organs mucfa modified from ttie 
tjiH.', that is from ttie leaf of vegetation. This ia simply what 
is meant hy the pro[K>sitioii that all these organs are transformed 
or nieUtmorphoaed k-avea. Wliat would have been leaves, if 
the development had gone on as s vegetative branch, have in 
the biosBom developed in other forms, adapted to other func- 
tions. Linn^ua expressed this idea, along with other more 
speculative conceiittoiis.dimljapprelieuik-d. by the phrase Vege- 
table Metamorphosis. Xot long aHerwards, this fecund idea 
of a common type, the leaf, of which the parts of the flower, 
Ac., were regarde<l as modifications, was more clearly and differ- 
eotly developed by a philosophical physiol<^st. Cas|)ar Frederic 
Wolff. Thirty j-ears later, it was again and wholly independ- 
ently developed by Goithe, in a long-neglected but now well- 
knowti essay, on the Metamorphosis of Plants. Twenty-threa 
years sfterwaTxls, similar ideas were sgain independently pro- 
pounded by DeCandolle, from a different theoretical jjoint of 
view ; and finally tlic investigation of phyllot«xy has completed 
the evidence of the morphological unity of foliaceous and floral 

1 The contribDIIon o( Linnteiu u on p. 301 of Ihe PhitiMuphia Boianica, 
1TS1 : and all that ii pertinent U in the fulliiwing propneitioni : — 

Plumulsm lemiois atspiuj terminat lot Hoa aut gemma. 

Prlncipiiun flonun et foXorum idem c>t. 

Prlncipium (^nunanim et folionim iilera cat. 

Gemma conitat foiiurum nidimcntls. 

Pcrianthium alt ex connatl* fotiorum nidimcntit. 

Hii duweruilrm, Prukpaii Hlantsrum. in Am(rn. Ac»4 rl. (ITflO), adiled 
nothing but otiwure gpeculalluni to the tonaei comparatively dear 

Kaipar Frieiirieh Wo]B'» contrihutlan ia in his Tlicoria Genirmtionia. 
imiinly concerning animala. poblisheil in 1760, and an enlarged and amended 
tditioii in 177*. He flnti cloarlj cnncelvei Ihe plant aa formwi of two ele- 
mcnla, atom and lof, but dt-vtlopa only the raorp'iolngy o1 tlw Uiier, and 
tuidcr the hypiitheila llist le»*« of rogWatlon become bud-scales or floral 
orfiuis, aa the case may be, through degenereicencc or diminutlun of tege- 
tstlve force, which la renewed in tlie bud or In thv H'ed. 

Johann Wolfgang Ofitlie'a Vi-nuch die Metamorphose der PfUniPii lu 
crkUmi waa pubiishcd lii 1700, In SO jmkm. For the iratislaliona and 
reproduL-tiona, acv Pritacl, llii-uuma. To Ihe French tnnalatioii by Siipet, 
with German text accompsnyinK (Stuttgart, ISJl), and alao to that of Cli. 
Hsrtvna ((Eur. lUat Nat. dc Gcelhc. Paris, 1837), are joiiiwl Ihe outlior'a inler^ 
estiiignoti-sandaneeilolcsodalcrperiods, down to 1831, The degencrcwent'e 
by diminution of vi^getailTe force with renewals by generation, propounded 
by Wolff. In Gotlw'a e»ay Ukcs the lorm o( tucceaitvo expansion and con- 
traction of organs. 

A. P. DeCandolU's Tb^orie £l(!meDlairc de Botaniqae appeared In 1813, 


305. It will be understood that metamorphosis^ as applied to 
leaves and the like, is a ligarative expression, adding nothing to 
our knowledge nor to clearness of expression, but rather liable 
to mislead. The substance of the doctrine is unity of type. Its 
proof and its value lie in the satisfactory explanation of the facts, 
all of which it co-ordinates readily into a consistent and simple 
system. As applied to the flower, two kinds of e\adence may 
be adduced, one from the normal, the other from teratohgical 
conditions of blossoms. The principal evidence of the first class 
is that supplied by 

306. Position and Transitions. As illustrated in the preced- 
ing chapter, the flower occupies the place of an ordinary bud or 
leaf-bud. Also the parts of the flower are arranged on the 
receptacle as leaves are arranged on the stem, ». e. they conform 
to phyllotaxy, as well in passing fVom leaves and bracts to the 
perianth, as in the position of the floral organs in respect to each 
other. This is partly shown in the preceding chapters, and 
is to be further illustrated. Sei)als, petals, stamens, and 
pistils are either in whorls or in spirals, and have nothing 
in their arrangements as to position which is not paralleled in 
the foliage. 

307. The evidence from transitions has to be gathered from a 
great variety of plants. Very commonly the change is abrupt 
from foliage to bracts, from bracts to calyx-leaves, from these to 
corolla-leaves, and from these to stamens. But instances abound 
in which every one of the intervals is bridged b}' transitions or 

a second edition 1810 ; a third (revised by Alphonse DeCandoUe), in 1844, 
is posthumous. The Organograpliic V^ijfetale, in which the morphology of 
the earlier work is developed, appeared in 1827. The leading idea is that 
of symmetry, of organs symmetrically disposed around an axis (the 
homology of foliar and floral organs not at first apprehended), but this 
symmetry disguised or deranged more or less by unions (solderings) of 
homogeneous or heterogeneous parts, by irregularities or inequalities of 
growth, by abortions, &c. 

The reason why the organs in question have a normal symmetrical dis- 
position on the vegetative and floral axes was not reached by DeCandoUe, 
nor was it perceived that the arrangement of leaves and of floral organs was 
identical. All this was the contribution of phyllotaxy, — a subject which 
was approached by Bonnet (an associate of DeCandollc's father), and first 
investigated by the late Karl Schimper and Alexander Braun, beginning 
about the year 1^20. 

It is interesting to know that Wolff's work was wholly tmknown to 
Gojthe in 1700, and that both Wolff's and GoDthe's were imknown to DeCan- 
doUe until after the publication of the second edition of the latter's The'orie 
Ele'mentaire, in 1810. When the Organographie appeared, the essay of Goethe 
had come to light ; and contemporary contributions to floral morphology by 
Petit-Thouars, K. Brown, Dunal, and Rocper, were adding their influence. 



intcmiediate forms The gradual transition from ordinary foli- 
age to bracts aucl bractltts is exceedingly common. In color 
and texture it is not rare to 
■ meet with bracts which vie 
I with, or indeed surpass, pet- 
f als tbemBelres in delicacy and 
^ brightness ; and in sucti cases 
they assume a principal office 
of flower-leaves, tliat of con- 
spicuous show for attraction. 
. Scarlet Sage, Painted-Cup 
f (Castilleia) , and tlie Poin- 
R settia, with other Euphorbias 
- of the conservatories, are ex- 
i aniples of this. In the flowers 
of Barberry, it is by a nearly 
arbitrary selection that bractlcts are distinguished iVom sepals ; 
in C'alycanthus, in many lands of Cactus, and in Nelumbiuni, 
the same is true 
as to bractlets, se- 
pals, and petals ; in 
Water-Lily (Nym- 
phiea, Fig. 3 IS), 
there is a gradual 
transition from the 
sepals througli the 
1 petals to etflmens; 
' in Lilies and most 
lily-like flowers, se- 
pals arc as brightly 
colored as petals, 
and common Ij' more 
or loss combined 
in with them. When 

the perianth-leaves arc of only one set, it is not at all by color or 
texture that this perianth can he assigned to calyx or to corolla. 
Normal transitions from a stamen to a pistil could not, in the 
nature of the case, bo expect*il. 

308. Terutologlcal TransltioDS and Changm. Teratology- is 
the study of monstrosities. These in the vegetable Icingdom 

no. 3i;. C*ctas-flo*cr IMunltlarls catpitdw), wllh bi 
Iiu>l»e fnlo atcU olhrr. 

Flu. 318. Seiie* exUblUng IranilliuD trom MpalB to omi 

In NyDiiituia odutaU. 


often elucidate the nature of organs.^ The commonest of these 
changes belong to what was termed by G(Btho retroffrade mtta- 
morphotit ; that is, to Tevertion from a higher to a lower fonn, as of 
an orgao proper to the eummit or centre of the floral axis into one 
which belongs lower down.' The most familiar of all such cases 
is that of the so-called double Jlower, better named in Latin ,^ 
plemu. In this, the essential organs, or a part of them, are 
changed into colored flower-leaves or petals. Most flowers are 
subject to this change under long cultivation (witness " double" 
roses, camellias, and buttercups), at least those with numerous 
stamens. It occasionally 
occurs in a state of nature. 
The stamens diminish as 
the supemumerarj- petals 
increase in number; and 
the various bodies that may 
be oflen obsen-cd, inter- 
mediate between perfect ^ 
stamens (if any remain) and \ 
the outer row of petals, — 
&om imperfect petals, with 
s small lamina tapering into 
a slender stalk, to those 
which bear a small distorted 
lamina on one side and a 
half-formed anther on the 
other, — plainly reveal the 
nature of tlie transformation 
that has taken place. Carried a step farther, the pistils likewise 
disappear, to be replaced by a rosette of petals, as in fully double 

' The lca<Ung Ircatlsea arc Moquin-Tandon'R Tc'ratalogie Vfg^tale, Paria, 
1841, and Maitcn, Vefcolahlc Ttratotnpy, London, publulicd (or the Bay 
Society. IStW. An earliiT publication desurvca pnrticular mention, vii. the 
theti* De Antholysi I'rotlromui, by Dr. George Engcimann, Frankfort on 
the M^n, 18-')2. 

* To these abnormal cliangei, the term mffnmorphinii \» obvioualy more 
applicable; for here wliat OTirlcntly should be atameni, pistils, &c., on the 
leatimooy of pnsllinn and tlie wlinle economy of the hloMom, actually ap- 
pear in the form of aome other organ : yet even here the chani^ is only in 
the nina formalieat : the ortfan naa not first formed aa a stamen, and then 
tTMiafonned into a petal or leaf. 

FIO, 319. A llnwer of the ennimon White Clover revertlnE to * leafy bnnch ; after 
Tarpln. Calyx wlLh tubo little dianKnl. but lubm bearing leafleU. Phtll aCalked; 
tba DTBry optn iown tlie inner edge, and the nuu^n* vt the pbUl^eat baulng leave* 



butlercupe.' In these the green hue of the centre of the roeette 
indicates a tenctencj- to retrograde a atop farther into eepals, or 
ijito a cluster of green leaves. This takes place in certain bloa- 
aoms of the Strawbcrr>', the Rose, &c. Such production of 
"green rosea," and the like, has l>eFn appropriately called 
ehU>ro$i*, or by Maatera c/iloranl/iy, fn>m the ehaiige to green. 
309. A monatroaity of the blossom of White Clover, long ago 
figured by Turpin (Fig- 319), is such a case of foliaceous rever- 
js, HI Bion, in irhioh even the ovules are implicated. 

Tlie iiDiwrfect leaves which take the place of 
tlie latter may be compared with the leafy 
tulla tvldeh form along the margins of a leaf 
of Bryophyllum, by which the plant ia often 
propngated. (Fig. 323.) 

310. The reversion of a simple pistil di- 
rectly to a leaf is aeea in the Double-Ootrering 
Cherry of cultivation (Fig. 320, 321), usually 
passing moreover, by prolification of the re- 

^ ceptacle, into a leafy branch. 

311. The reversion of pistils to stamens is 
1 rarer, but has been obscn'ed in a good number 

" of instances, in Chiies, in the llorsiTadish. 

in Gentians and Hyacinths, and in some Willows. In the 
latter, the opposite transformation, of stamens to caqx-ls. is 
very common, and curious grades between the two are met with 
almost every spring. So also in the common Honseleek, and 
in perennial Larkspurs. Certain apple-trees are known, both 
in the United Slates and Euroi>e, in which, while the petals are 
changed into the ap[>Garance of minute green sepals, the outer 
stamens are converted into carpels, these supernumerary and 
in the ftruil suiwriwsed to the five normal carpeb.* In Poppies, 
many of the innermost stamens arc occasionally transformed 
into as many small and stalked simple pistils, surrounding the 
base of the large compound one. 

■ It muit not tie concluJtil that the gujxmunicrar]'' potals in ■)! such om.-* 
Bie reverted •tamen*. or ■tamcnsaudpUtilti, Soruu mre inilanct-s of shnornuil 
pleiolaij, ie. al tlie production of one or mart addiijoiial ranks of petnJi 
(bcttiT de»CTTinK the o»roe of JmiUe Jioutr). with or witliout revereiiffi al 
cucntiBi organi to flower lea vea. 

* These tree* arc popularly euppoaod to bear fruit williout tiiiwaoniitii;: 
the reverted green pctali bdng >o inconspicuoiu that Uic flower i« un- 

YIQ.XM.3H. arMnt«*M)h>ni 
one itlll nbtnrlnK, try lU pKrdBl Invo 
carpsl, ilie either * noAll bqt nrll-tnnDni Mar. 

FIO. SSI. Loaf or [mUH of Drjioptiyllum. devi'l oping plantlcle almg tbo margliu. 



312. Another line of teratological evidence is Aimished by 
prolifieaiion. The parts of the flower ore, by the doctrine, 
botnol(^u8 vith leaves, and no leaf ever terminates an axis. 

Normally, in fact, the . 

axis is never prolonged -.A 

beyond the flower, but /i\i«i-' 
abnonnatly it may be. ":a^ v 
It may resume vegeta- \\^ 
tive growth as a termi- \\y 
nal growing bud, either f^--^ " 
from between the pistils 
after the whole flower is 
formed, or at an earlier 
period, usurping the . 
central part of 
flower. Thus, when a 
^ rose is borne on a pe- 

duncle rising from the centre of a rose, which is 
not Tciy unusual, or a leafy stem from the top 
of a pear (Fig. 323), the flower was probably 
complete before tlic monstrous growth set in. In 
Fig. 324, the reversion to foliaecoua growth toolt 
efiiK;! after the stamens but before the pistils ** 
were formed. In rose-buds out of roses, the terminal proliferous 
shoot takes at once the fomi of a peduncle ; in the shoot Stoxa 
the pear, that of a leafy stem. 

313. A^rain. axillary- buds are normally formed in the axil of 
leaves. No such branching is known in a normal flower. But 
in rare monstrosities a bud (mostly 
flower-bud) makes its appearance in th 
axil of a i>otal or of a stamen ; and it 
may be clearly inferred that the oigan ' 
(not itself asillan) from the axil of/ 
which a bud develops is a leaf or i 
homologuc. Fig. 325 exhibits a clear 
case of the kind, a flower in the axil 
of each jietal of Celastrus scandcns. Flowers, or pednncolate 
clusters of flowers, from the axil of petals of garden Finks are 
sometimes seen. A long-i>cdunculatc flower from the axil of a 

nan pMr. prnloiigBl into ■ Icaff brunch : Tma Bonnet. 

Lo mstKinriTphniikii nl ■ iawn at Iha PraxliwIlB of tbs gkntsni, 

>riTnrt1rDltiua; ui[nleniod(i>loDg>l«dJuMaboTatha>Uinaiia, 

of raliw BIttei m wwt (Galwtnu nandnii) pradndnf otlwr 

FIO. 323. A tn' 

PIO, K4. Bfln 

IWiin LInillfV'a T)» 

■nrl hnrinR k irbn 

Fia. a». A fl 

174 THE PLOWEtt. 

stamon of a. species of Wntcr-Lily (Nymphiea Lotus) is figi 
ami descrilM-d by Dr. Masters.' 

31J. In the application of morithological ideas to the elucida- 
tion of Uic tlower. notUiug should Im: assnmed in regard to it 
nhieli lias not its proper countcrpurt and cxetDplar in the leaves 
and axis of vegetation. 

Section II. Floral Stmmetbt. 

315. The parts of a flower are symmetricaUy arranged around 
its axis.* Even when this symmetry is incomplete or ini{>erfeet, it 
is still almost always discernible ; and the particular numerical 
plan of the blosBom may be observed or ascertained in some of 
the organs. 

SIC. Adopting the doctrine that the parts of the flower are 
homologous witli leaves, the syinmetry is a consequence of the 
phyllotuxy. It is symmetry around an axis, not the bilateral 
Bynimctry which jtrevails in the animal kingdom. For parts of 
a flower disposed in a continuous spiral (which mostly occurs 
.yben they are numerous) , the arrangement is that of some order 
F'Uiis kind of phyllotaxy, which distributes the parts equably 
> BUpcrposed ranks. (237.) The much commoner case of 

' The fullMt cniinieratioD aod dlicuwiion of the very Tariou* kindj of 
•hnurnisl ilrucluro snd dcrlatioas in plants la lu be found In ihe TiTBtology 
uf 1)1, MhbIiti, nlnri! rcfem-d to. Many U'chnical (I'nns arc here brought 
Into twc, width n«'il not be lipremcntiontd, except the following, which relate 
(Urectl; to Burol loelamorphoiiia, 

Phi/ttody (called I'kiflloiBor/Jii/ by Hoirrn, Frondaceiux by Engelmsnn) i» 
thfi condition whcn-iji true leave* are eubilituted for lome other organi ; 
I. r., where oilier orgMns are netamorplioseil into green Ivatei. Tlierv la 
phyllwly uf jiiitila, uvulea, fllaments, anther, petala, sepals. &e. 

Stpaiadi/, where otlier organa aiaume the appearance of green aopala. 

Prtaiodg, wlirre they auunie the appearance of pelala, m normally In 
PlDckneyaMid Calf caphyllum. in whieh one ealyx-lobcenlargee and becumit 
p«tal41kc, and ahnornially in Priniroica where all Ihc calyx-lobca iniitalo 
lobMot the corolla I thia baa been lermcd CaigcardKtmy) ; alto of tlie »laniea« 
of eunimon " doublu flowen." 

Statniiiodf. where other organ* develop into atamcni. Caaea of thia aa 
affecting piallla aiv referred lo ahore : rarely aepala and petal* are lo affected. 

PtUUlodg, where other organ* develop into pblila, which moat rarely 
bappcn* except with the alatnena, aa above mentioned. 

* II la ataled that Conva de Serra (who publjahcd botanical and other 
paper* In London, Parli, and Philadelpliia during the fint twenty yean of 
the century, but who knew tarmorc than he published) waa the flrat botaniit 
10 ln«tat on the ayniinf try of the flower. It waa flrai made prominent by De 
Taiidoile, In the Ttii^rie fCKinentaire, and elaborated in detail by A. St. 
Hilaire in lii* Horpbologiv Vfg^talc. 


equal number of parts in a cycle, and the cycles alternating 
with each other, is simply that of verticiDate ph3'llotaxy. (234.) 
In either case, the members of the successive circles (or of 
closed spirals as the case may be) will be equal in number ; that 
is, the flower will be isomerous, 

317. A Symmetrical Flower is one in which the members of all 
the cycles (whorls or seeming whorls) are of the same number.^ 
In nature, the s}'mmetiy is of all degrees : it is most commonly 
complete and perfect as to the floral envelopes when it is not 
so as respects the essential organs. The general rule is that 
the successive C3cles alternate, as is the nature of true whorls. 
But the supeq)osition of successive parts is not incompatible 
with S3'mmetr3* of the blossom, although it is a departure from 
the ordinary condition, assumed by botanists as the t}^)^ An 
isomerous flower (meaning one with an equal number of mem- 
bers of all organs) is the same as symmetrical, if the reference be 
to the number in the circles, rather than to the total number of 
organs of each kind. 

318. A Regular Flower is one which is symmetrical in respect 
to the form of the members of each circle, whatever be their 
number ; t. e., with the members of each circle all alike in shape. 

319. These two kinds of s}Tnmetiy or regularity, with their 
opposites or departures from symmetry, need to be practically 
distinguished in succinct language. For the terminology, it is 
best to retain the earlier use, generally well established in phjix)- 
graph}', as above defined. 

320. A Complete Flower is one which comprises aU four or- 
gans, viz. cal^'x, corolla, stamens, pistil. 

^ This is not only the definition " generaUy appUed in English text-books/' 
but that introduced by DeCandolle, adopted by St. Hilaire, and foUowed at 
least by the French botanists generally. The innovating German definition, 
of a recent date, is tliat a symmetrical flower is one " that can be vertically 
divided into two halves each of which is an exact reflex image of the other." 
But such have immediately to be distinguished into " flowers which can be 
divided in this manner by only one plane," which Sachs terms " timpljf 
symmetrical or monosymmetrical" and those which can be symmetrically 
divided by two or more planes, "doubly aymmetrical or pdyaymmetrical^** 
as the case may be. Now both these forms have a more expressive and 
older terminology, adopted by Eichler, viz. : — 

Zyf/omorphous, for flowers, or other structures, which can be bisected in 
one plane, and only one, into similar halves {median xygomorphous, when 
this is a median or antero-posterior plane, as it most commonly is ; trans- 
verse zygomorphousy when the plane of section is transverse or at right 
angles to the median, as in Dicentra) ; 

Actinomorphous for flowers, &c., which can be bisected Id two or more 
planes into similar halves. 



331. Vnmerial irroand-plan. Many fiowora are numerically 
indetinite in some or most of their kinds of membera. as Banun- 
cnliis, Magnolia, and the Rose for stamens and pistils, Nym- 
plisua for all but perha|)3 tiie sepals, many Caetacem for all but 
the pistil, and Calyenntlius for nil four components. But more 
commonly eacb flower ia coustracted npou a definite numerical 
ground-plan ; and the number U usually Ion. Seldom, if ever, 
ie it reduced to unity in a hermaphrodite blossom (even Hii>- 
puris, with a single stamen and a single pistil, is not on un- 
equivocal case) , and probably never in a complete one. But there 
are such extremely simphfied flowers among those of a single sex. 
In Mouocotjledona, the almost universal number is three, some- 
times two; in ordinary Dicotyledons, five prevails ; four and two 
are not uncommon ; three is occasional ; and higher numbers are 
not wanting, as twelve or more in llouaeleeks. 

S22. To designate tlic particular plan, such familiar terms of 
Latin derivation as binary, ternary, quaternary, quinary, senary, 
&c., a're sometimes employed, denoting that the parts of the 
flower are in twos, threes, fours, fives, or si.xes. More technical 
and preeiac t«rms, equivalent to these, are comjKisGd of the Greek 
numerals prefixed to the word meaning parts or members, as 

Manomcrout, for the case of a flower of one memln^r of each ; 

Dimerous, of two. or on the plan of two memlters of each ; 

TVi'meroiM. of three, or on the plan of three membera ; 

Tetramerotu, of four, or on the quatemarj- plan ; 

Penlamerous, of five, or on the quinary plan ; 

Hexamerovt} oT six, or on the plan of six members to each 
circle. But, in Jlonocotyledons, so-called hexamerous blossoms 
are really trimerous, the sixes being double sets of three. 

323. Fattom Flowers. These should be tymmetrirnl, regular. 
enmpUu in all the parts and without ex- 
cess or complication of these, and with- 
out any of the cohesions or adhesions 
^ which may obscure the ti-pe. or render it 
j leas expressive of the idea that a flower 
J eonsists of a series of cireles or spirals 
of modified leaves crowded on a short 
** * axis. Wherefore the illustration Fig. 307. 

with ita diagram Fig. 308, may serve as a jiattem pentamerous 
or quinarj- flower; and Fig, 326, with its diagram. Fig. 327, 

' ThrBemBvbe«liorllywritt«il-meroua, 2-nieniug, !^iiiOToni,auJ<ooniip 
to lamrroug \dttai*en»ii). IS-mcTona {dodrcamrrmit), 4c 

FJ«. XX. Pxrtu iir ■ ■finiDsEilcBl trlnisroul flnwur (TJIIXI rnoKO**): a. iMjx; 
6 cnmllaicMMDciii; il i>I*i[1l 3£7. Dlncrim aribouiDe. 




as a pattern trimerous or ternary flower; these being simply 
isomerous, and of one circle of each kind. And the whole 

relation of the parts, viewed as 
modified leaves on the common 
axis, ma}' be exhibited in such a 
diagram of a pattern isostemonous 
5-merous flower as that displayed 
in Fig. 328. 

324. Diplostemonoiis l^jpe. The 
foregoing patterns are selected 
upon the idea of the greatest 
simplicity consistent with com- 
pleteness. But extended observation leads to 
the conclusion that the typical flower in nature 
has two series of stamens, as it has two series in 
the perianth; that is, 
as many stamens as 
petals and sepals taken 
together.^ As - the 
petals alternate with 
the sepals, so the first 
series of stamens al- 
ternates with the pet- 
als, the second series 
of stamens alternates with the first, and the pistils or carpels 
when of the same number alternate with these. Thus the outer 
series of stamens and the carpels normally stand before (are 


* This yiew of the symmetry of the flower was first taken by Brown 
(Obs. PI. Oudney, in Denham and Clapperton Trav. 1826, reprinted in Ray 
Soc. ed. of Collccte<l Works, i. 203). It is true that Brown declares the 
same of the pistils ; but that is not made out. The eyidcncc of this* doctrine 
is to be gathered from a large and varied induction ; from the general pres- 
ence of the two sets of stamens, and no more, in petaloideous Monocotyle- 
dons ; the unaltered position of the carpels (before the sepals) when the 
inner set of stamens is wanting, as in the Iris Family ; the rery common 
appearance in haplostemonous flowers among the Dicotyledons of vestigea 
of a second series, or of bodies which may be so interpreted. The androecinm 
or the blossom is said to be 

Ttostemmous or Haplostemonous when the stamens arc of one series, equal 
in number to that of the ground-plan of the blossom ; 

Diplostemonous, when there are two series, or double this number. 

FIO. 328. Ideal plan of a plant, with the simple stem terminated by a 83rrametrlcal 
pentameroas flower ; the different sets of organs separateil to some distance from each 
other, to show the relative sltnatinn of the parts. One of each, namely, a, a sepal, b, 
a petal, c, a stamen, and d, a pistil, also shown, enlarged. 

FIO. 329. A pentameroas dlplostemonous flower of Sedom. 




superpoacd to) the aepala, and the stamens of the inner series 
stand before the petulfl ; as in the tliagram, Fig. 331. ^ 

325, Flowyrs which completely 
escmplil^- their tj-pe or sjinmetry 
are rare, but most exhibit it more 
or less. Eacli natural order or 
group exhibits its own particular 
floral type, or modification of the 
common type.' Some of these 
modi Ilea tiona do not at all affect 
the symraetrj- or obscure the plan 
of the flower, except by combina- 
tions which render the phjlhne 
character of the floral envelopes and carpets 
less apparent, such combiualious being of 
rare occurrence in foliage. Others gravely 
interfere with floral ajinmetry, sometimes to 
I aucU degree that the true plan of the blossom 
i to be ascertained only througli extended 
coniparisona with the flowers of other plants 
of the same order or tribe, or of rekted 
orders. The symmetrj' of the blossom finds its explanation in 
the laws which govern the arrangement of leaves on the axis ; 
that is, in pliyllotaxy. The deviations from sj-mraetn,- and from 
tjincal simplicity have to be explained, and in the first instance 

t reference nnd Ihc Rvotdance of circuuilocution, mme 
writer* lerra ihe etamcns which aiv before the p«tali tpipeialom, those before 
the tcpaU tfimpalout ; but. as this prefix means upon, it is better to restrict 
these temu to cases of ndnation of atameiu to these respecUre parts of the 
perianth, and to itislinguieh ■• 

Aniipelalimi, those Btnmeiui which stand before petals, whetlier ailnate or 
free, and 

Aniittpal<mM, those which stand before sepals. — These terms we find hate 
already' been employed in this way by Dr. A. Ditluon {in Soeniann, Jour. 
Bot iv. 2TC), with the addiilon of a third, \\t. 

Parapflaioiu, for stamens which stand at each aide of a petal, yet not 
neuessnrity before a sepal, at in many Rotacen. 

■ ITieie particular types, with their modi flea tions, are set forth in the 
ehanieitri or distinguishing msrlis of the orders, tribes, genera, ftc The 
best ^nerally aTailable lliustnitions uf ordinal type* are in I« Maoat and 
Decaisnc's TraJte Gt^niTal de Boianlque, and in Hcmker'a English edition 
and revision, entitled A General System of Botany, Descriptive and Analyti- 
cal, London, 1ST3. The best ninrpl'olo^cal presentation is in Eichler's 
DlrithcndinRTamme, &c. (Flower Diat^rams, Constructed and llliutraU'd). 
Leipzig, lt5T5. 

FIO. 330. Opgnsd tovret of TdlUam ai 

, 331, Dlagnm of the SB 


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 t^-pe. 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: Irregularitt. 

4. Non-appearance of some parts which are supposed in the 
tjTpe : Abortion or Suppression. 

5. Non-alternation of the members of contiguous circles: 
Anteposition or SuPERPOsrrioN. 

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 fVom assumed pattern are seldom single ; 
possil)!^' all may coexist in the same blossom. Several of them 
occur even in that one of the orders, the Crassulaceae, which 
most obWously exhibits the normal tj'pe throughout. 

328. Thus, Sedum (Fig. 329), with two circles of stamens, 
being taken as the true t}'pe (324), Crassula (Fig. 307) wants 
the circle of stamens before the petals ; Tilhea (Fig. 32G) is the 
same, but with the members S3Tnmetrically reduced from five to 
three ; Rhodiola loses all the stamens b}' 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 

middle ; Penthonim (Fig. 335, 336) has ita Ave carpels coales- 
tTcnt almost to the top. and usually loses its petals by abortion ; 
In GrammanUies and Cotyledon (Fig. 332-334), the sepals are 
cualescent into a cup and the petals into a dcei>er one, out of 

which the stamens appear to ariae, these being adoatc to the 
corolla. Symmetrical increase in the number of members of 
each circle is no proper deviation from tj-jw, at leaat in this 
Ikmily (in which flowers on the same plant sometimes vary Trom 
a-merous to 4-raerou8 and G-merous) : and in Sempervivum {to 
which Houseleek belongs) these members are always more than 
five and sometimes as many as twenty in each circle. 

ReouLAR UmoK ov SIMILAR Parts. 

329. Coftlescence. 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 gjnflecium and corolla. Fig. 
471— 4 7G further illustrate it in the corolU, and in various degrees 
up to entire union ; and Fig. 483-488 illustrate it in the andns- 
ciura. The technical terms which coalescence calls for, and 
which are needfd in botanical description, may be found under 
the account of the particular ot^n, and in the Glossary, Such 
growing tt^ctlicr of eontigiions members in the blossom is strictly 
paralleled by conn ate- perfoliate leaves of ordinary foliage (212, 
Fig. 215), where it more commonly occnrs in upper leaves, and 
in bracts, which are still nearer the flower. 

330. It should now be hardly necessarj- to explain that the 
terms coakaeence, cohesion, union, and the corresponding phrases 

PI0.331. Flonr of GrumiBiiUiei- 3Aa Flnmr '>f it Colj-IMon . SM.ThewroIlft 
ilil npFO •hoiTliig tlia two nm ot •tamcui ln«rlal on It. 33S. Ths flto pliULs uf 
BulhoruiD, united 3X. A cruw ■ o cUua of tliB tmna. 


in the next paragraph, do not mean that the parts were once sepa- 
rate and have since united. That is true onl}' of certain cases. The 
union is mostly congenital, equall}* 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 undinded 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 inseparate. 
But the common language of moq^hology 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 '^ insepa- 
rate " base of a calyx or corolla, consists of a certain number of 
these. It is no contradiction to this view that they developed 
in union.^ 

§ 3. Union of dissimilar or successive Farts. 

33 1 . Adnation 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 tlie lan^nuigc of descriptive 
botany in tliis regard, consistency would require its total reconstruction, with 
the abolition of all such terras as clefts 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 aa 
connate, adnate, coalescent, &c., which imply and suggest the fundamental fact 
in the structure of phxnogamous 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 thallua 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 diversiflcation 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. 

r ont of SBOtfaeri - 

182 T 

]ippiu«Dt gnwing of cue put on or o 
the rorolU oui *if tho cnI^-x. the stamras out of the eoroD^ ov 
aU of ibpm oat of the pistil. Thb 'lisgatws the real or^io 
ot the floral or^ns bam the tecepUcle or axia. to Hiccesaive 
series, one within or shore the otbn. Onguis in this cooditioD 
are also ami rigiitly said to be Otmmalt (bom onitMl) ; hot, as 
this tena is eijnaltr applicable to tfaa coalnccitee of members of 
the same circle, the word Aiiitalt ia preferable, as ajifilving to 
the present case oaly. Adnation is hetcrc^eneaos organic oo- 
besioti or adhesion: coa- 
tesceoc? is homogeneous 
o^ieston or aniou. 

332. AdnatitHi occnra io 
■ ery various degrtvs, and 
' atTocts either some or all the 
organs of the floncr. Its 
consideratioQ introiluces into 
terminology several peculiar 
terms, which may here be 
defined in advance. Three 
of them, introduced and 
prominently employed by 
Jussieu, depend upon the 
degree of adnntion, or the 
absence of it, viz. : — 

ffjfpogynoui (literally be* 

neatli pistil) . applied to parU 

which are imerted (i. e. are 

borne) on the receptacle of 

/*^^~~- i*Ti'''L5 (^ 1 "** *'""'^''' "8 '" ^'8- 336. 

/^ ^NtTwHCTT ^ M-v This is the absence of 

~ adnation, or Uie condition 

which corresponds with tlie 

unmodified tj-pe. 

PerfffynoHt (around the 
pistil) implies an adnation 
■■ which carries np Ibe inser- 

tion of parts (which alwajs means aj)|«ireiit origin or place of 
attachment) to some dietanco above or away from the reccji- 

■nnii Fisi. tbnirinc ihi nmBk] or 

OI |XUI> UpUO IM EOTBI or tWDplUle. 

MDilonnf ■tBrnraraftlieCliBn}', la ahawUiB perleynona liiaetUon, 
OKI Jl, or tbB pelkla uul Mimenii. 

Inr iMUnn of Ibi Siiwar ot tha PimUne, ■bawlug an ailniitlon tf kll 


taclc, SO commonly placing this insertion aronnd instead of 

beneath the pistil ; whence the name. The perig}'ny may be, 

as the fi^ircs show, merely 

the ailnatioD of petals and 

stamens to calyx, the calyx 

remaining hjiragynons, as in .' 

Fig. 337 ; or else titc ndno- f 

tion of the calyx, involving V 

the other organs, to the lower 

IMirt of the ovnry, as in Fig. 

338, or tip to the summit of 

the ovary, wliile the ]>ctals *" 

and stamens are adnate still (brther to the calyx, as in Fig. 389. 

The latter [msses into what is called 

Epigt/noat (on the pistil), where tbe adnatiOD is complete to 
* the ver>- top of the ovarj-, and none beyond it, as in Fig. 340, 
341. Yet here the [>art8 so termed are not really on the ovarj-, 
except where an epigynous disk (494) actuaUy surmounts it. 

333. Adnation brings some other tenns into use in botanical 
descriptions, es]M>cialIy those of superior and iufcrior. In this 
connection, these words (in Latin taking the form ofiupentt and 
Jn/erut) denote the position in repect to each other of ovary and 
floral envelopes, — not the morpholf^cal, but the apparent posi- 
tion or placo of origin. Thus, in Fig. 336 and in 337, the calyx 
is inferior, or in other words the ovary mperior. Here real and 
ap]>arcnt origin agree, this being the normal condition, which 
is otlierwise expressed by ea.ving that the parts are/r<«, t. e. ftee 
fVom all adnation of one to the other. But, in Fig. 339-341 , the 

FIO. 339. SlmiUr Hctlon of ■ flnwar oT Hk*Uloni. ibowlng compteU ■dnMIOD to 
theoDinnilt ofEbenTury BTiilor the other p«U bcjnnU. 

FKl. 310. Venlciil wvtlon oT n Cnnberrr-aoirer. anil Ml, at Homr of Anllk 
nuitkaull«. with h-ciUoiI e[>1i[]'nni»iDKitk>nof cal]ri, ei^olU, viil ■Umeni ; Uwnljx 
of the laiur conpletsif cautulhlkted wlUi tb« luclhiH of tba antj, or IM Umb 


ovary is aaid to be inferior and the calj^ ti^erior, the calyx 
and other parts, in conaequence of the adoation of its lower part, 
seaming to rise from the siunmit of the ovary. 

334. Adnation of floral envelopes to pistil rarely extends 
beyond the ovary; yet, in apecies of Iria hani^ a tube to 
the perianth, this tube is commonly adnatc for most of its length 
to the style. But when the caljTC has its tul>e or portion with 
united sepals prolonged, the petals and the stamens are usually 
adnate morc or less to it, t. e. are inserted on the ealffx. 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. 

835. 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 Cherrj' (Fig. 337), stamens 
as adnate to base of corolla in Fig. 334. and a long way farther 
in Phlox, dec. That the calyx is simUarly adnate to the ovaiy 
is nearly demonstrable in certain cases. 

336. But, OS the lower portion of a pear is undoubtedly recep- 
tacle, or rather the cnlai^cd extremity 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 Cercus 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 eal^'x-tubc adnatc to an inferior 
ovary is partly or wholly, a hollowed receptacle (in the manner 
of a Fig-fniit) ; that is, a cup-shaped or goblet-shai>ed develoi> 
mcnt of the base of the floral axis. This would bring the case 
under S 7. (82G, 4y5.) 


337. ImsnlBrlt;, or inequality in form or in union of mem- 
bers of a circh', is extremely common, either with or without 
numerical ayramrtrj-i One or two examples may sufllce. 

838. Irregular flowers with sjTnmetrical perfection, except in 
the gyntecium. are well seen in the Pea Family, to which belongs 
the kind of corolla called Papillonoeeom, from some imagined 
reflemblance to a butterlly. (Fig. 342-341.) This flower is 
fi-merous throughout, has the tbil complement of stamens (10, or 
two sets), but the gjnojcium reduced to a single simple pistil. 
The striking irregularity is in the corolla, the i^etals of which 
bear distinguishing names : Ihe posterior and lai^r one, exter- 
nal in the bud, is the Vbxillcu or Stakdabd (Fig. 844, a) ; 


the two lateral next and under the staDdard, Al£ or WiKOS (b) ; 
the two anterior, covered by the wings and partly cohering to 

form a prow-shaped body (c), the Cabina or Keel. The calyx 
is slightly irregular by nneqnal union, the two upper sepak 
united higher than the 
other three. The sta- 
mens are much more 
Coalescent,butivith an 
irregularity, nine com- 
bined by the lower part 
of their filaments, and 
one (the posterior) 
separate. (Fig. 345.) 

839. The plan and "■ « 

floral sj-mmetry in the Locust- blossom and 
its relatives are little ohsciired 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 arc distinct or unconnected. Only 
the calyx is more irregular, by the union of the two posterior 
sepals almost to the tip. (Fig. 348.) 

FIO. 312. IMignm of flowerortbe L«ait,RabltitkPMad*eacl*: a. axlioflnflDna- 
eenre; b. btact; fliTt ciri'le of B. caljx: Hto remiilnliiK piece*, comlln neit sntlieri, 
tre a Klngledmpla rl"')' 3*3 Front ileir df Lociut-flower 

FKI. MS. AnrlrOH^Iu 
imo Ufa Laplne, All te 

iT Ibo Locafit. nlno siAmtni co&lesoent. 
lamenti onnleHant helnir <nu ■ fIiwiI t 
irolla of B^pUila ftutnlli. He. Suns 
■boirliig Um dUUiict »'■"■"•■ uid Upottlm itjla. 

a40. Butii 


Lupine-blossom, of equally near rektiocuhip, 
a casual obsorvpr might fail to rcct^ize Ihe very same tj-jie, 
although rtiagiiisHi only by cohesions. For while the two pos- 
terior sepals are unit£(l to the tip on one side of the blossom, the 
three others are ainiilarly umt«d into 
one iMKly on tbe anterior side, giving 
the npi>carance of two sepals instead 
of five : iu tlie corolla, tbe two keel- 
petals are more atrietly united into a 
slender scythe-shnped or sickle-shaped 
body ; so that the petals might with 
tlie unwary pass for four: in the 
andnpcium, the coalescence includes all 
ten stamens (Fig. 346), wtijch is an 
apjiroach to regularity, 
fd^^^rt T ^^ — N 341. The o-meroua symmetrj- of the 

^"^"^^ [J jj^"—- ^ Violet-blossom is complete- until the 

^' JVl '^s, gjHttcium is reached (but with only 

/ I A one circle of stamens) ; the mai n irregu- 

W/nVi larity of the periouth ia in tbe aulerior 

\_^ petal, with its nectaiiferious sac at base 

"■ (Fig. 349-351} ; the two stamens near^ 

est tliis send into the sac curious appendages, which the other 

three do not possess ; the gjiioicium is composed of tbi-ee car- 

S t>els coalescent into one compound ovaty in s 

■ hereafter explainetl. In Antirrhinum 

^^^ and Linaria (Fig. 480, 4S1), there is a similac 

^\?\ iiTcgularity accompanying coalescence of the 

I ', / petals, the anterior one being extended at base 

^' ' into a nectariferous sac or hollow spur,' The 

flower of a Lolx^lia (Fig. 488) has Die same 

numerical plan and sjinmetrj- as that of Viola 

(eiicopt that the gjatecium is dimerous) : but 

■e adnate l)olow and coalescent alwve, and the 

corolla is irregular through unequal coalescence of the five jwtals, 

ami the absence of coalescence down one sitle. 

' Pelobia ie a name ){iven "by Linnsiu to an occuional monuairoait; o( 
theie flowen (imiiattil in 6undry(ithm},iairhich the base of ever]' petal, or 
answmn); part of the corolla, la prolonged downward into a lac or apnr. 
Tbi- lac u, morphologically considered, a departure from normal ri'gularily ; 
in the montter, aynimetrical ri'gularil; iarcstorvd li; Ibc development of four 
more aacs. 

no. MS. Finww »( VIM* MgltlaUi. 3,W. IM mymXt an'l prtali •llsplarcal. 3S1. 
Diagnunara Vlolei-btmnir, trnni Elcl^ler, vlih bn<:I<<raii1itim<llng leaf (bvlnirl. a pair 
ur biiictloW (lateral ), abd axis lu wLloh tlio luUuuiUug leafbvliingidlKivuat ivntiiiorj. 

the members s 




342. Abortion or Suppression are somewhat s^-nonymous terms 
to denote the obliteration or rather non-appearance of organs 
which belong to the plan of the blossom. Abortion is apphed 
particularly and more properly to partial obliteration, as where 
a stamen is reiluced 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 irregularit}'. 

343. Of parts of a Circle. Among papilionaceous flowers 
(338) , different si)ecies of Erji-hrina have all the petals but one 
(the vexillum. Fig. 344, a) much reduced in 
size, in some concealed in the calyx, and in 
everj' 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, 35;5.) 
This one is evidently' the vexillum, both by 
position and sha^K; ; and the 5-merous t3'i)e, 
also the particular ty^Ki of the family, are still 
discernible in the five notches of the cal^'x, 
the ten stamens, &c. In a related genus, 
Parryella, even this last petal is wanting, and 
the andrcecium is straight, all irregularity thus 
disai)i>eanng through suppression. 

344. Delphinium or Larkspur and Aconite or Monkshood 
furnish good examples of flowers in which irregularit}' 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 upi)ermo6t and largest, is pro- 
longed posteriorly into a long and hollow spur. Within these, 
and alternate with them as far as they go, are the petals, only 


FIG. 352. St&menB and pUUl of Amorpha frutioota. 3S3. An enUre flower of the 


ftinr in nmnbcr, and those of two shapes, the two npiM^ 
hnring long spare which are received into the spur of the upper 
sepal; the tiro lateral ones having a small but broad bUde 

raised on a fltalk-like claw ; and the place which tbo ffflh and 
lower pctaUhoiild occupy (marked in the ground-plan. Fig. 356, 

FIO. SM. Flower uf « tArlu|>ur, ass. The Dve wpMl* (oaler clrctH) uid tbs fain 
tiBUI>lliiDatdRleldl>tila)«d. XC Omunil-pten ortlwc*l)'i and comlla. 

FIO. 35T. Flower u( tn Aconite or Monkibouil. 3U. Tlie lite niatln ami the two 
BDall aiid flurloiul]' (batial |«Ml> <Il*i>lared : alea ifan iisBieiii ami pUUli In iho centra. 
au. GroD»I-p1aiiar(lNc«lixaadcarolla: tbo dcUed line*, a* lu Fl^. IW, reproMnUiv 


bj a short dotted line) is vacant, this petal being suppressed, 
thereby rendering the blossom uns^Tnmetrical. 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-sha|)ed or hood-like body; 
three of the petals are wanting altogether (their 
places are shown b}' 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 they 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 extraordinary' 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 irregularity of the perianth, especially of the corolla, 
characterizes a series of natural orders with coalescent petals. ^ 
These flowers are all on the 5-merous plan (except that the 
gynoecium is 2-merous), but with corolla, and not rarely the 
cal^'x, irregular through unequal union in what is called the htla- 
hiate or two-lipped manner. The greater union is alwaj's median, 
or anterior and posterior, and two of the coalescent members form 
one lip, three the other. The two ix)sterior petals form the 
upper lip, the anterior and two lateral form the lower lip of the 
corolla ; in the cal^'x, 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. 361, 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 

* These natural orders in which this occurs, or tends to occur, are the 
Scrophulariaceae (Snapdragon, Pentstemon, Mimulus, &c.), Orobanchaceae 
(Beech-drops), Lentibulaceae (Bladderwort), GesneracesB (Gloxinia), Big- 
noniacesB (Trumpet Creeper, Catalpa), Pedaliacese (Martynia), Acanthaces, 
Labiatse (Salvia, Stachys), &c. 

FIQ. 360. A petal (nectary) of an Aconite, much enlarged. 



filamenti — a monstrosiU', indeeil, but the monstrosity la here 
a return to noniml symmetry. The two stamens nearest the 
suppressed or abortive one generally 
' share in the tenrtciicy to aboiliou, as 
is shown by their leaser length or 
smaller anthers : in the flower of 
Catal|>a, these two also are either im- 
[•erfect or reduced to mere vestigea 
(as in Fig. 363) : in very many other 
plants of these families, even these 
vestiges ar« not seen, and so the fire 
stamens are by abortion or complete 
suppression reduced to two. 

34G. Suppression in the gjna>ciam 
to a numlier leaa than the numerical 
plan of the flower (aa shown in the 
perianth) is of more common occur- 
rence than the typical number, and 
the retluction is comparatively con- 
stant throughout the genus or order. 
A papilionaceous or otiier leguminous 
flower with more than one or with all 
I five pistils is esceedinglj rare, and 
except in one pentacarpellar}" genus 
is a monstrosity. Suppression of 
' the interior is more common than 
of exterior organs. Want of room 
in the bud may partially explain 

347. Snppresslon of whole Clrolos. 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 

IneompUte^ in which any oneor more of the four kinds of organB 
is wanting, whatever these may be ; 

ApetaUitu, when the corolla or inner perianth is wanting : 
MonoMamydeoiig, where the jwrianth ia simple instead of 

PIO. Ml. Oin.11* of n..nHi» i>Drpur» Uld 

whlrh U» Bftb ihonlit '-vavy ln.U«H-1 \.y n rn. 

■ Ulitapcii.wltli 

no. Ma. CurullAutCauljA UlUcpva, wilt 

1 two porfwC itao 




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 ; 

Achlamt/deausy 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 ; 

Monacious (of one household), when stamens and pistils oc- 
cupy different flowers on the same plant ; 

Dicecious (of two households), when they occupy diflTerent 
flowers on diflerent plants ; 

Polygamous^ when the same species bears both unisexual and 
bisexual or hermaphrodite flowers. This maj' occur in various 
ways, from the greater or less abortion of either sex, either on 
the same or on separate individual plants ; — as Moncsdously or 
Diactously Polyyamous^ according to the tendenc}' to become either 
monoecious or dioecious. Recently Darwin has well distinguished 
the case of 

Gyno-dicBcioHs^ where the flowers on separate individuals are 
some hermaphrodite and some female, but none male onlj' ; and 
Andro'diaecious^ 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 onl}' 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 



Clematis and Anemone, Fig. 364) , and the corolla is sometimes 
grL'eiiiali or leftMike, the only real difference between the two is 
that the calvx rt'presenta the outer 
nnd the corolla the inner aeries. 
Even this distinctioa becomea arbi- 
trary when the periauth coDsists of 
thj'ee or four circles, or of a less 
definite number of spirally arranged 

349. Yet the only perianth ob\-i- 
~fl ously ])resent may be corolla, as when 

m the calyx has its tube wholly adnale 

to the ovarj- and Its honler or lobes obsolete or wanting.' Arnlin 
nudieaulis (Fig. 341) is an Instance, likewise many Umbelliferie, 
some species of Fetlia or Valeriauella, the fertile flowers of Nyssa, 
and those Compositie wbicL have no pappus. For Pappl'S, 
the name originally given to thistle-down and the like, answers 
to the Iwrder or lobes of a calyx attenuated and depauperated 
down to mere fibres, bristles, or liairs. The name is ex- 
tended to other and less oblitcrste^l forms. (644, Fig. 631-633.) 
When the obliteration is complete, as in Maj-weed (Fig, G30), 
in some spet'ies of Coreopsis, &e,, the corolla seems to be simply 
continuous with the apex of the ovarj'. A comparison with 
related forms reveals the real state of things.* 

3^0. 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 ovarj' (epigynoue) 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 

' In the Haven at the two common species of Prickly Aili [Zmlhoxy- 
luml of ihe Ailuilic Unilcd Slntei, one hn* » double, ilic oihcr n <iiiglo 
perianth (na Bhown in Gray, Gen. Hluilr, il. 148. t. 16G) : llic iMwirion of the 
■iruncna ^veta presuinpt ton that themitung eircleof tlic Ulterisihecaljx; 
yet it may bo olhemiK explained. In SanuUcOB thcK arc aoine groanda 
for iuipcclinti that the ilmple perianth, although opposite the ((ameni. la 
corolla : anil the foliaceoiu aepal-lobes nf the female flowen of Bucklcya 
would contirm ihig, if thoie are true aepali rather than adnate bracts. 

' In the pappna of Compoaitv, every gradation U teen between undoubted 
calyt. recoimioible aa anch by alructure aa well aa posillon. ami lUaphanotia 
M-»le». I>ri»Tlca. and more hairi. wholly " trichome* " a» to itmcture. although 
in the place of " pliyllome* " and repreienlinK llieni. 

no. MH Ftnwgr of Anemnns pntniri'anlea: ■pnUliiu*. U>* falrr pctaloid. 

riO.SU Aclilam)'<I«HuacHrBaILliard'»-liUI(SaurDrui<wni ' 




trimcrous) flowers of Saururus, Fig. S65. But acblamydeovt 
blossonta are usually still ftirther reduced to a single sex. 

3'i2. Suppression of one cinle of stamens is of ver}' common 
occuiTcneo. It is seen in different species of Flax ; which have 
mostly 5-mcrouB perfectly symmetrical and complete flowers with 
one set of stamens abortive. In some species (as in Fig. 307), 

vestiges of the missing circle of stamens are conspicuous in 
the form of altortive filaments, interposed between the perfect 
stamens ; in others, these rudiments are inconsptcuons or even 
altt^c titer wanting. 

3o'd. Suppressed Andrtedan or Ornieclam. This occurs with- 
out or along with suppression in the perianth. In cases of the 
fonncr, vestiges of 
the aborted organs 
often remain to sig- 
nil^- the exact nature 
of the loss. yei>a- 
ration of the sc.ics 
[monacious, di'aci- 
out, <&c.) is tlie re- 
sult of such suppres- 
sion. In Menis[x!r- 
this is accompanied 
by an actual doub- 
ling of both calyx 
and corolla. The 
dicBcious flowers of 
Srailax are similarly 

complete, except by ^ "" 

the alwrtion of one sex, but the caljx and corolla are single. 

no. Xe. Flnwe 

niimarFUx. 36T. AndrtBclnn 



370-373) I 

3.'t4. Comblanl ntth snitpressloD of Perianth. Tbia, whU-li is 
fmiiid in most anicntnffoiia or calkin- bearing trees, iu some willi 

Sliniliiil suppression of perianth, is well illuslriileil 
iu Willows. Ilie flovfcrs of wUich nre all achliiinydtoat 
find diaciuvi. (347.) The little scale (glund or nec- 
tary) at tlic inside of each blossom might be bu[>- 
posed to represent, a perinntli, reduced to a single 
piec« : but an e^ctcnded pomparisuu of forms i-efere 
it rather to the receptacle. Willow- blossoms ( Kig. 
e crowded in catkins, each one in the axil of a brait : 
the staminate flowers consist of a few stamens merely, in this 
species of only two> and the pistillate of a pistil merely. In 
SaUx purpurea, the mule flower seems to be a sinple st.imen 
(Fig, 374) ; but it consists of two stamens, uniu-d into one 
bo<ly. Here extreme 
suppression is ac- 
companied with CO- 
aleei.'euce of the 
existing mem Iters. 
■— \ \r-;i' / ir ^.^ ^'*^' ^*'" ™*''^ 

fjJ V, ^T \Iy^^ simplified flowers, 

but more dillicult 
to eonipifhend, are 
those of Euphorbia, 
or Spurge. These 
are in fact montcci- 
ous ; and the female 
flower is a pistil, the 
male is n stamen. 
The pistillate flower 
(of Ihree carjwla, 
their ovaries united 
into one fhree-lobed 
comjxinnd ovarj) 
surmoimta a slender peduncle which lenninntes each branch of the 
flowerintr plant. (Fig. 375.) From around the liase of this Jie- 
ilnnclc rise other smaller and shorter peduncles, each IVom the 
axil of a slender bract, and surmounted by & single stamen, 
which reprcscnla a male flower. (Fig. 376, 377.) Tina nmlteUikc 

Kh; 371 A vp>r*la •tonlTiBte nirvBtiif Sallx ptirpniwi. witli iImj ■lunieni ««•- 

'"''"'>' ii'ltih'to* Mwl •yna*n*iliHiB|. «» uiu tpimtr lUu ■ dnolo one. 

I'll, :r,: Klnvtnrlug bnuwh at KnphnrliU curnllnl*. JT«. Calyi-llli* IntrJuef* 

.ihi.f.'.i :. iiL:ri.ul~', a1iuwlu(tba>tiiinlnM>n<>i(eraarnunil«frt«111Man(i»n'ln). ATT. A 

."niii.iihii.] l.y tlic •nllury iinmi^ii. e. 319. PliIU In tTa\t, col omm. ihonlnj Ihn 
Ihi™ iiiiu-rwclfl cun""!" "f "hlfli It I" compowd. 


flower-cluster is surrounded and at first enclosed by an involucre 
in the fomi of a cup, which imitates a calyx ; and the lobes of this 
cup (the free tips of the calyx-leaves) in the pi*esent 8i>ecies are 
bright white, so that the}* exactl}' imitate petals. Here, then, is 
a whole cluster of extremely simplified flowere, taking on the 
guise of and practically* behaving like a single flower, the invo- 
lucre serving as cal^'x and corolla ; the one-stamened male flowers 
collcctivel}' imitating the andrccciura 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. Sappression of both Andr<ee{iim and Oynoecinm. This 
occurs in what ai*e termed Neutral Flotcers (347), such as are 
conspicuous at the margin of the cjTiies of Hydrangea (Fig. 293) 
and of Viburnum lantanoides and Opulus, also at the mai*gin of 
the head of flowers of Sunflower, Coreopsis (Fig. 287, 288), and 
the like. In these and most other instances, the perianth of 
which onl}' the flower consists is much larger and more showy 
than in the accompanying perfect flowers : in fact, their w^hole 
utility to the plant, so far as known, is in this conspieuousness. 
No plant normally bears neutral flowers onl}- ; 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. Occasionall}' flowers become sterile 
and neutral 1 >y mere depauperation and abortion of i>erianth as well 
as of essential organs, as in certain Grasses; but such are 
mostl}* vestiges of flowers rather than neutral blossoms. 


357. Anteposltion or Saperpositloii is the opposition of succes- 
sive (or api)arently successive) whorls which normally alternate. 
This result is brought about in different waj's, some of which are 
obvious, while of some the explanation is hyix)thetical. 

358. In the first place, there are cases of seeming anteposl- 
tion, which are explained awaj' on insixjction. In a tulip, lil^', 
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 
G-merous, but 3-merous : there is a calyx of three sepals, colored 
and mostl}- shaped like the three petals, which alternate with 
those and are clearly anterior in the bud ; next, three stamens 
alternate with the f>etals or inner circle of the ixirianth ; then 
the three stamens of the inner circle, alternating with the preced- 
ing, necessarily are opposite the three petals, as the first three are 


0])i>o8ite the sepals. Tbesc organs altogether arc in four whorls 
or three, not in two of six raemlwrs ; and the iiistil at the centre, 
of three <.'OiuhiDe<l members, is the fifth and Bnal whorl. 

3o9. The Barberry family exhibits a similar secmiitg nnte- 
position, which is more striking on acconnt of a multiplication 
of the members of the i>erianth. The calyx is of six sepals in > 
two circles, the corolla of six petals in two circles, the Btamcna 
equally six ; and so each petal has a stamen before and a sepal 
behind it. But, when proi«rly >'jewed as a trimeraus flower with 
double circles of seimls and [totals as well as of stamens, all i 
symmetrical and normal. J>[eiiis[]ennum in the related Moonseed 
family is in the same case, bnt the flower is tetramerous, as seen 
in Fig. 369 : in the mate blossom this is obscured in the andros 
cium (Fig. 3G8) by a multiplication oi' the stamens.' The 
same thing occurs in the i»rianth and bracts of certain Clustaoese, 
in which the members counted as in fours are superposed, and 
in some of which the double dunerous arrangement with apparent 
ant«position extends through the corolla ; while, in other closely 
related flowers, the corolla changes to simplytetramerous and to 
alternation with the preueding four sepals. This passes, in the 
same family and in the allied Temstmemiaceie, into 

3(10. Snperpoiltion bf Nplrab, as where five ]M<tals arc Bat«- 
Ijosed to five seiinls, by an evident continuation of iwntastichoua 
pliyllotasy ; an<l the stnmcn-clusters of Gordonia Lasiantlioa 
arc probably in tliis way brought before the petals,' The flower 
of Camellia is continuously on the spiral plan up to the gjTMB- 
cium : luit uimn one which, from the bracts onward, rises f^om the 
) to till' If auil I onlcr or higher, throwing the petals of the rosette 
in a r\illHloublc flower into numerous more or less conspicuous 
verticiil ranks. 

3(11. Antepoflllion In the Andrfficlam. It is in the andriccium 
that ri'ul anteiHisitiun is most common, and also most tliOicult to 
account for ii|M>n any one principle. Doubtless it comes to pass 
in more than one way. This condition is chiefly noticed when 
tile stamens aiv dellnil^ in number, and mainly in isostemonous 
and diplostcmonous flowers. (324.) 

3Gi. With IjMHlomonr. Vitis (Fig. 379-381). also Rhnmnus 
(Fig. -11^, -tif)), and the whole Grape and Buckthorn families of 

' Li Columljino (Aiiiiilrtrio). niulliptiuRlion nf iht' tlmoc-ns in BUix^eatiirclj' 
alrvrnHtlng O-meruue wliurli wmilitrlf lirinniK Ihe niiilrnviuni inlo Icn miiks : 
su. wIr-ii thi>ae ctimMii In iluublc Huwcrs niv InmafoniiiHl itito bo]luW'a]iurT(.>d 
pctali, thrtc'arc tct one into another in Ivn vcrilcnl ranlu. 

= Gen. niuicr. ii. 1 140. Bdi the ptlnli alleniale with th<? ippBla in the 
onKimry nmnncr of ihc Hoirer, though Ihvir (tronH quincunvial imbricKtliin 

• UBgl-'H 

III ml H 




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 
the receptacle, alternate with and inside 
the stamens : there is no good reason to 
suppose that the}' answer to a second 
row of stamens. All isostemonous For- 
tulacaceoe 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 
gamo[>etalous corolla, such anteposition 
is universal in Plumbaginacese, Primu- 
laceae, the related Myrsinaccae, and in 
most Sapotaceae, in the latter usually 
with some complications. 

3G3. The earliest and the most obvious explanatioh of the 
anomal}' is that of the suppression of an outer circle of stamens, 
and to tliis view recent morphologists are returning.* 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 rudimentar}' organs exactly in the place 
of the wanting circle of stamens, which ma}' well be sterile fila- 
ments. In the allied order Sapotaceae, while Chrj'sophyllum 
has in these resixjcts just the stmcture of Primulaceae, and 
Sideroxylon that of Samolus, Isonandra Gutta (the Gutta-percha 
plant) has a circle of well-formed stamens in place of the sterile 
nidiments of the preceding ; that is, alternate with the petals, 

1 Eichlcr, Bluthendiagramme, passim, and in preface to Part II. xviii., 
relating chiefly to obdiplostemony. The principal opposing view is that of 
St. Ililaire, 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 
androDcial 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. 38L Diagram of the flower. 



cofu pic ting the syrutnctrr of tho lilossom nnrl the nomuil alterna- 
tion of its members. This explanatioii of the aiiteiK>sitioii of a 
single circle of stnmeiis is the more reailily receiveil, because it 
well afoonls with the idea here a(loptc<], tliat the niidnrcium 
of a typical flower slioultl consist of two circles of stamens. 
(324. ) The only serious objections to this explanation rise out of 
the diftculty of applying it to analogous antejioHition when both 
uireles are present. 

36i, For Dipl<it!emong, the cowlition of two circles of sta- 
mens, eacli of the same unmlwr as the petals, is ako itself vcrj' 
commonly attended hy anteiusition. In nontml or Direct 
Diiilotlrmoni/, — that which answers to the floral tvpe com- 
pletely. — the autisepulous stamens (324, note) are tlie outtr 
and the antiiietaloiis the inner serice, and the carpels when 
isomeroiis alternate with the latter and np])osc the sepab ; the 
alternation of whorls is thei'cforc comjilete, as in the diagram, 
Fig. 382. Such stamens, however, maj' actually occupy a siugtn 
line or coalesce into a 
S' S tulic, without derange- 

uieut of the ly|>e. But 
commonly it occurs 
1 llmt the aiitipeiiiloui sta- 
19 arc more or less 
/ exterior in insertion, and 
then the carpels, when 
uerous, are alternate 
"" "° with the inner and anti- 

supnlous stamens, and thcrefoi-e opposite the ijetals, as in tlio 
dia^am, Fig. 383. This arrangement takes the name of 
Ofnliflla$iemoHff. In it the normal alternation of successive 
whorls is intcriiipted. so as to produce anteposition, 

3ii5, With Obdlplosleraonjr. This condition prevails, niore or 
less eviilently, in Erieacea', Oeraniaceie, ZygopliyllaceiE. Riilaeete, 
SaxilVngnccrc, Crussnlaceas Onagmcew, &c. (hut in some of these 
with exceptions of direct diplostemony) ; also, aeeompanied 
by a peculiar multiplication of members (380), in Malvacete, 
RtcrculiaoeiB. and Tiliaccce. The exi>lanation is dilDcult. The 
hypotheses may lie reduced to three, neither of which is quite 
satisfhctor)'. There is, Orst, the hypothesis of St. Hilairc, ait- 
plied to this as to the preceding case (to Rbamnna, Vitis, &c.), 
that these exterior antipelalons stamens l>olong to the c^orolUne 
whorl; In otlicr words, timl the petal and (he stamen Iwfore it 


(whether ndnate to or fVee from it) answer to one leaf which 
has developed iato two oi^ana b}' a dcduplieatioti (372) takiog 
place transversely. This makes the inner and 
anti^palous stamens the thii-d floral eirclc 
or the only tnily andnreial one, and sjra- 
metrically alteniate with the petals oe the one 
hand and the carpels on the other. The 
second hypothesis conceives that there is a 
whorl su|ipreased between these anttpetalous 
stumeus and the corolla : this, ideally restored, ** 

gives symmetric succession and alteniatiou to all the aucceeding 
whorls. The five glands in a U cranium-flower, alternate with and 
next suceeedingthe)>etaU (Fig. 384), were plausibly supposed to 
represent this missing whorl, which according to Urauii should be 
an inner corolla ; according to Others rather a primary circle of 
stamens. The thirtl is the recent hypothesis of Celakowsky, 
which Kichlcr 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 hyi>othe3i3 are, first to account for this dis- 
placement, and then for the ante|)osition of the carpels to the 
assumed inner ataiocns in the great majority of these cases.* 

> In Ihe flrst pan of tlie niiitlieniliagrammc, Eiclilet ini;Uned to Ihe Drat 
hypotlicsia, lliat of St. llilairc (now very n]ui:h nbandoned on account of 
tlic feeble evidcni« that tliciv ii any such tiling as tranaTt^rK or median 
tliorisiu); in (lie apcond, lie discards tliii in favor of Ci'lakowsky's view 
(pubtislicil in ItegunBliurg Flor.t, 1876). A» to members which are morplio. 
loflicatly interior becoming exterior by outward displaL-enient, Kicliler uitei 
Ihe slaniinwlia or sterile stamen-clusters ot Famassia (Fig. 400, 401), and 
the correspunilinK aiitipetaluus stamens of Limnantliei, a* clearly interior 
in Ihe early Hiiwer-liuil, but exterior at a later period ; slates Ihftt the vascu- 
lar bundles which enter these stamens Kencnilj; are either inner as respccis 
those of the episcpaluus stamens or in line with them ; thai in sonic eases 
(as in many CHryo]iliyllaceK) the real insertion of the stamens is that uf 
direct dtpluntemony, while the upper part of Iheir filaments and the anthers 
arc external ti> the epi«epali)us series; lliat in most families with obdiplos- 
teniony examples of direct diplostemnny oecur,and still more cases with both 
Blamineat circles inserted in the same line ; and that, as a rule, the epiecp- 
alou« stamens are either later or not earlier formed than the epiiwialous. 
As to tlic position of the carpels before anilpelaloua stamens and petals, 
Celakowsky suj^iests that this may result from the outward recession of 
thiwe stamens affonlinfc more room there, while in the normal ease Iho 
greater space is over the cpiscpalons stamens. And, indeed, exceptions 
to the prevalent position are not uncommon both in direct diplostemony 
Fill, .114. DlAcrain lirnM-sertliiiil of tlie flower nTGcnknliim maciiliitum, eililbltliii 
tbr TBialive txwltinii nf puriM. ami tlie synimetrlral alEernailun of ilrcles, i. e. seialr, 
petals. gtoeiilaU Lvilltis callot Kiaiuls, autli«taluus scsniBiis, anttsepaloai M»-~'"- 

200 TlIK riXJWBE. 

^I'li, Tho caac of ftUimciis in a cluster bcrorc tlic petals is a 
iiuin{ilii-iiLl(fti ur t-itlii^r of tlie forfgoiiig witfa a pccniiar kind of 
nitilti|)Iicutloii, iktiiijdkutioii or chorisie. (372.) 

(7. I.N( 

SC7. Aiifmeiitatltfii iti the nuint>cr of floral mcmlicTS !a odg 
(}f lUe cuniiiKiluiit iiii^lillctitloiiH of the lype. It occurs in two 
wiij'n: Int. \iy iin iiicrcawl nniiibt-r uf circles or turns of siiirals 
111 the Huwfr, wUif^h U Jtfyular AfuHiplicaliou ; 2d, by the pn>- 
duotiun of two or tlir(« or of timuy organs in the normal place 
of one, ChoriMtt or fJudupUeatiun. The first rloes not alter the 
normal Bjrametry of the blossom, nitliough it may render it dif- 
fleult or iin[K(Sbible to trat:e or dfmonHtratc it. The Hccond 
»p[iBrenUy dlHliirbM, or al least disguises, floral symme^-. 
Kitlier may lie dr^niiK, or of a (instant and comparativdy 
small iJimilKT; or indefnite, nhen too nuraerone for ready 
counting, or ineotistant. aa the higher numbers are apt to he. 

.tliN. Ib^olar Hultlpllcatlon, or Augmenlaiion of Jlural cirdet 
or tpirah, may iilTcet any or all the foui- organs, but most com- 
monly the andiieeium. When the perianth is nmeti increased 
in the iinmlKT of its inemliers, the ilistiiietion bctwe«^ii calyx and 
corolia. or even between bracta and corolla, is apt to disappear, 
as in iiioHt Cac'taeeous flowers (Fig. S17), Nclumbium, Calycan- 
thns, &.i: In tlicsc and similar cases, the members of the jKrianth 
are prone to take a spiral instead of cyclic arrangement ; and this 

tnil In ub(li|il»tL-imin]'. Mang with the luck of clear nnslogy to supixirt 
Bt, llllalrp'a lir|>olliralB of tranivtric diHlupMvRllnn, the similar orii-nUIiun 
of the TRai^uUr biindlo* In the petal and the suiiiun bcfon; It niuat. tu 
CvUkatrtky Indnti. lie gixKl pvldi'nce tlioi tlietc represent inilcpcnilent 
IciTo. snii not aiiiiFnioM^I porllnn* uf one. 

Tlie riMlii oliji-i-lloii to the ucotkI hypotticaU (llial of & sapprcMed 
drcio oataiik' uf iliu nnttpt'Oloui mtamcnal ii thnt thi* mining circle, 
wlii-tlicr of peula nr (taniL'na, la not actually tnct with in an; nearlj r» 
Uli!)] forini (for In Mnnmnia (he flttrcn atamrna are otlirnriie explained) ; 
alio ttial there are traiiRiiiona. a* ahovo tncnlfimeil, Iwlween obdlploiiemonj 
and direct illploatvmony, Tu Itrnun's theory ihiit the ifland* liehind the 
aiilliepaloiu atainena In true Gcraniacnr ana wit to auppreaacd phylla, 
Eicliler ubjpcia that thcav arc prcacnt behind pII ten atamcns in Oxalidcv; 
aUo that all are wanting when tlic offlee of nectar-aeerclion, which tliey cub- 
•erve, la underukpn by aoinn other part of tlie flowtr, oa by the calyx-*pur 
In Pi'lar^nltini an'l Truixwtluin. 'Ilic first nbjeciiuQ ia forcible : ilie tecoud 
mixn morphulii^i-'Nl eunalilcraiiuns wlili functional, and ia inconcluaivc. 
AlKiniic organs, pmcrvcd for their utility a> nrctarlca, might totally die- 
■I'Pear whiui rendered uaeleai bjr s different provltlon for tlie tame function. 


is even more true of greatly multiplied stamens and pistils, as 
in Magnolia and Liriodendron, most AnonaeeiB, 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 calj'x and corolla bj* the 
production of one additional whorl of each, and the seeming 
ante(X)sition which comes of it when the andrcecium remains 
simply diplostemonous (in the manner Of the Berberidaceae, 
Menispermaceae, &c., 359) has already been explained. 

370. Similar increase to two whorls affecting the corolla onl}' 
characterizes Anonaceie, Magnoliaceae, Papaveraceae, and Fiuna- 
riacea?. In all but the last order, this is accompanied b}' indefi- 
nitely multiplied stamens, and mostl}- b}' an increased number of 
caqxjls. In Fumariacea*, which has dimerous flowers, there is a 
diminution by the suppression in most cases of half the normal 
andrcecium, and also an augmentation of the other half by 
chorisis. (372.) 

371. ParapetaloQs Maltiplication. Under this head may be 
described an anomalous arrangement of augmented stamens 
which prevails in the order Rosaceae, but is not peculiar to it.^ 
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 i>etals ; but the space between two stamens before each petal 
is mostl}' rather wider than in the pair taken the other wa}-. 
The next case in order, as in 15-stamened Hawthorns, and 
constanth* in Nuttallia, adds to the above a simple interior circle 
of five stamens, one directl}' 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 successiveh' the later.* 

1 It was first clearly dcscrilxMl by Dr. A. Dickson, in Trans. Bot. Soc. 
Edinb. viii. 408. and Seenmnn's Jour. Bot. iv. 473 (1860). He introduced the 
term, fxiraftetaloux, which is characteristic of it in its elementary form (254, 
note) : it is particularly illustrate<1 by Kichler, in BlUthendiagramme, ii. 
4ft5-510. 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. 

^ Accordingly, the whole is probably to be explained by 80^ 


Sli. ChorUls or DednpiicatioD. Itoth tlieae tonus, ond the 
idcna which they rleiioto, oiigiiialeH with Dunal, liut wi-n.' flrst 
ex|Ktiiinlpcl by Moijain-TaniJoii,' The first wonl is (Jreck for a 
scparutitig or scii&ratuiri. Thu second is a translation of Diinal's 
French word diduuUemcnt (Itlvrally undoubling), the ambiguity 
of which, aud of tha original pruscntation of the case, lung 
retanled the right appreheiiwon of the subject. DirempUon has 
bt'fn suggested (lij- St. Ilihtirc) as a proper term. The mean- 
ing similly i». the division of that wliich is morphulo^cally 
one oigaii into two or more (a di^'ision wliich is of course 
congeuittti), so tliat two or more organs occupy tlie position of 
one. As thus used, chorisis is restricted, or nearly so, to the 
hoRiologues of leaves in the flower, aud mainly to stamens and 
(»r|}els ; the division or splitting up of a petal or a bcjjdI, when it 
occurs, Ix'ing expivssed in tlie phrases whicli are appliyil to leaves. 
Yet a coniixjund loaf, es|>ecially one of the pulmat« type, is 
a good type of chorisis, the several blades of a ^-oinpound leaf 
answering to the single blade of a simple leaf. It has been ob- 
jected against tlie terms ebovisis and deduplicntion that they 
assnme the dinslon of that which has never been united ; but 
so equally does the established tenuinology of foliage. A di- 
vided leaf has never been entire. 

373. Chorisis is complete when the parts conccme<l are dis- 
tinct or separate to the very insertion, as in the stamen -clusters of 
Ilj-pcrienm. The foliar form of this wonld Iw represented by 

tlon n( the uugtiK'nMUnt) of cirelci. Dieksun't hjixithcib, ihnl the two, 
Ihrw, or five aiBmcn* wliich nre more of Jess In fate of pnvh pelnl are nil 
dedujiliealiuiia uf tluil pc'tnl, would voitii> to bv iiotknl under the nexl liead, 
Itul it nuiy bi> diiniiucd at ooiv. Yet tliut the pain in (he nuter cinrle 
rcpresi'nl I'Beh an anlliepalous itanirn, divided by c)iori*ii (BOinetiaiF* 
Ineomplvivljr) and mueli wparatcd, la not improbable. The oilier lenabie 
explanation (whieh may be biiniinniied wlili ihe lail) ii Ihnl the outer 
circle al ilaaiena here riglitly roiwialt uf ten nieml^cn, reapeclifcly altemnl- 
\ng with tlie Be|i<il> awl p«-IBla taken aa a vrhulo. Tlili makes Uiem para- 
pvtaluiia, anil at Ilie same time bring! them under Ilofmeiitter't general law 
that new oritan* originate over intt-rral* of thoae preceding, in lhi*ea*eDVi.-r 
the ten pcrtanllf Intervals direcily. Ii at»o aeconU with Harto^a elueidaliun 
n( lh« acceMorj parta In the lluwer of Suputaei'n ( in Triroen't J»nr. Bot. 1878). 
The inner ctreles arc there (ometinm rvmerout after the primitive type, 
•aroptirow* lU-merous in rvtiuUr alternation to the preccdinm drclca. 

I Hoquin-Tandon, Kual dca lJi<doablemonf. £c., Monijietliur, JVSO; Cmr 
•id^lioiM snr les Irregularity de In Camlle. &c., [n Ann. Sei. Nat. xxvU. 
237, 1839: T<<mTolni{ie Veg<<lnl«. 3-17. DnnnI, Kssal siit Its Vaednit^i, 
IBlfl. elled liy Moqiiin (aome pagv« printeil, liut never publiihedl : Cansid^- 
ratloiia »ar U Nature et l« llappurt* de quelquit-mu dos Urgnne* de la 
Kleiir. IKH). The next butaniit to develop It wiu St. Uilaire, Murphotogie 
Vtfpitale, INl. 


atich sessile palmatc]y compound leaves as those of some species 
of Aspalatlitis, It is incomplete when division does not extend 
to the base; as in Kig. 387, 303, Compare, as a <C^ff/) 
proximate homologuc of this, a petal of Mignonette, "^J^ 
Fig. 385. But pro|)er ehorisis requires that the ^/?^ 
sujiomumerar}' organs should be developed like W '. 'j 

unto the original organ which is thus multiplied, or \^ 

should complete their sjmnntrj, whatever it be. ■ ms 

374. St. Ililaire distinguished two kinds of deduplicatiou; viz.. 
coUatertd when the memliers stand side by side, and parallel 
when an oi^n becomes double or mtdtiple aiitero-posteriorly. 
The latter, sometimes calknl rertical, and sometimes trantrerte, 
is lictter nanicnl median ehorisis. The collateral is the origi- 
nal and typical ehorisis. Most botanists incline to restrict the 
name to tins, and to give some other explanation and name to 
the meiliaii form of augmentation. But some cases, such as 
those of Titia and .Spnnnannia, are clearly of the same nature 
as the collateral, and nwy l»e a disguised form of it ; there are 
others which may be explain(.>d in accordance with it ; and there 
are such transitions between some of these and coronal out- 
growths that the tcnu ehorisis is most conveniently made to 
comprise augmentation oi' doubling in either plane. Distinct 
antoposition. however, may l>e explained in other waj-s, (3J7,) 

37^. Typical or Collateral Cborisk, in which the members, 
together answering to one leaf, normally stand side by side, 
occurs in many families of plants, and 
in a variety of forms. A few are here 

370. Elodes Vii'ginica (a common I 
marshplant of the Hypericum family), 
like most of its near relatives, has its 
calyx and corolla on the pliin of five- 
its Htamens and caqx'Is on the plan of *" ™ 
three, as is shown in the diagram. Fig. 380. This makes a break 
in the symmetry Ix'tween the corolla and the Stamens; but all 
within is in rcgidar alternation when the tliR-e stamens of each 
cluster are counted as one as their union at base into a plmlanx 
(Fig. 387) may suggest. These phalanges alternate with the 
three car|K'ls, and therefore stand where single stamens l)elong. 
The three cons|)icuous green projections, which in a general way 

FlO, 3M. A [flal nr MiKnonello (K««e>U odiinita). witb manjr jiutol blade, 
FIO. 3>Ji. Dimn-Bin of (Irtwrr of Elwlen Viririn!.'*. wiili tliree phalavj/fi at bUthfiu 



ai'e called ff/ands, altemati? with the phalanges, and bo are taken 
U> represent the outer circle of stamens. The nior])hologist 
aci-ordiiigly sees in the glnnds the homologuca or representatives 
of the outt'r series of stamens, reduced to three bj abortion, and 
in the three stamen-clusters only the three alternating stamens 
of the inner series, trebletl by ehorisis, and this chori^is incom- 
plete, because it has not quite divided the filament into three. 
In Hypericum, the ghinds are completely suppressed, eath pha- 
lanx is almost or ({utte divided into a cluster, either of about 
three stamens each, as iu H. .Sarotbra. or of a few more (in H. 
tnutilum and H, Canadcnse), or of an indefinite number, as in 
tlie common Ht. Julinswoits. Then in some otlier sj^eeies (as 
in our H. pyramidatum) the caq)els and the stamcn-clustcrs rise 
to five, realizing complete pentamcrous symmetrj', except that 
the almost numberless stamens all belong to the one inner circle. 

Morpholugically, thej' 
in most sj>ecies three) 
decompound and ses- 
sile or almost sessile 
leaves. The indefinitely 
numerous stamens of 
Rieinus are similuily 
incwased fVom five tv 
compound ramiflcation, 
377. Fumariaeeic.tlii 
Fumitory family, n]ii\ 
Airnish the next i\lii>- 
tration. The Bower is 
on the plan of tno 
(dimerous) throughout. 

mparable to the le&flets of Ave (or 


Taking Dicentra to show it. there is first a pair of small and 
8ca]e-sha]>ed sepals, not unlike the pair of bmctlcts on the 

Fia, 3m. I>lf*iiiniCMulUrla(nuWhnuui'iBrBocti«),»iriip 

WTuml ftatn Ui« ■liiiular bu1b(r>>rTnoil of Tltr rnlnrinl buoi of IV 
Inn'M', oT nttarnJ dni. •Iinirinc ston Otr pair at liracttaU ni> iIm 
»ini \.utt illiiiUfml. »iid Sin. Iniwr ivinls pluwl nlmia 3W. 



pedicel below (Fig. 389, 390), and normally alternate with them : 
alternate with these is a pair of larjje ix»tals, decplj* saccate or 
spurred bc»low ; alternate with these, a pair of smaller petals 
with spoon-shaiXHl tips which cohere at the si\x^x (the corolla 
therefore of two circ*les as in the Related Poppy family) ; alternate 
with these, two phalanges or united stamen-clusters, of three 
stamens each ; alternate with these is nothing, for the setrond 
set of stamens is wanting : ulteninte with this vacancy is a pair 
of carpi'ls wholly comhinwl into a comi)Ound 2-merons pistil. 
The statement itself explains the morphology. The three sta- 
mens of each phalanx stand in the place of a stamen, and arc 
the divisions of one. In l>i(*entra the memlKTs of the phalanx 
are almost separate; in Adluniia (Fig. 39.')) and Corydalis the 
undi\ided iilament reaches almost up to the anthers. The middle 
anther of the phalanx is normal, or two-celleil ; the lateral 
anthers are one-celle<K as if halvwl.* 

' Eiclilcr ailoptfl tliiin iiiterpri*tati<»ii (prop()$4*d in Gray, Gvn. Illustr. 
i. 118), and apftlics it to the iTuciai instance uf Hy]Hronni. In the tlowor 
of thu Old Wfirld genus, tliero are four nppiirently t^iniple and complete 
stamens, one before each petal : the simplest interpn'tatitm wouhl lie that 
which the factM apiH*ar to present, viz that Ix^tli dinu^rous circles of stamens 
are complete and nonnal. Hut Kichler — in view of the early development 
and the double vascular bundles uf the stanieuii iM'fore the inner i>etaU, and 
some occaitional Kli^lit disjunction of their anther-i'clls — considers that 
the interior utamen-i'ircie i!« wanting here, no less than in the other ^*nera 
of the onler; that what here takes its ]>Ince U'fore each inner petal is a 
stamen composi'd of the adjacent lateral memU'r of the phalanx, con^reni- 
tnlly 8evere<l from the ^rou]* to which it Ulon^rs and soldered into one fila- 
ment, bearing; the two oni'-cvlled anthers so bmught to^'tlier as to imitate a 
normal twn-celU-d anther. The or^ano^i'uy of the blossom is thou^rht to 
favor this hypothesis ; and it certainly favors the view hen* adoptcMl of 
the composition of the tliree-inemlx'ri'd phalanx of the family p.'nerally. 
If this interpretation of IlyiK'coum seems far-fetched, it is no more so than 
its exact counterpart, through which IM'andolIe, Lindley. and others explain 
the case uf the rest of tlie family. Starting; with that ^mius as the simple 
type, they conceive tliat the stamen opp(»sed to each inner petal is each 
pe%'ere<] into two. ami that these half-stamens attached to the sides of the two 
intact stamens, thus pnMluciii^ the ])halan^es by coalescvnce. 

A gcHxl empirical conception of the formation, fn>m a pintle leaf, of three 
stamens in Fumariaceas or two in Crucift-ne, is affonknl by the ])etals of 
Hypecouni. as illustratefl by Kichler. The outer jH'tals are slightly thn-e- 
lobe<l from the afK'X ; the inner are deeply so and nam>wer. The mem- 
bers of the next circU* in the family generally are just such three-h)lKtl 
bodies, the tip of each IoIk* transfonned into an antlier. There is an a]>- 
parent wmgruity in the pniduction by the symmetrical middle lobe of a 
symmetrical two-colled anther, and of a one-celled anther by each unsym- 
metrical latenil loU* or stipule-like portion. A fuller development of these 
sides of the leaf, and non-<levelopment of the middle portion (somewhat 
after the analogy of I^thyrus Aphaca, Fig. 219), with anther-formation, 
would convert the leaf into a pair of stamens. 


S'S. The obvious rtlHtionsliip of Cnicirenc to Fiimariaoeie, 
\; |(wiiliarity of having the Iwo carpels 
siile by siilu instvad uf furu anil oft 
(mi'dian), uiiri Ihc fharacU-ristic 
fliiomnly which the andnrciiim pre- 
iunts (i. ft tlic tetimljDani.v). wouhl 
j iiive reaaou to expect that its proli- 
8 might be solved by cliorisia. 
ludrtKl. the doctriDO waa applied to 
tills, long before its B[>plication lu 
tJic other order. Beginaitig at the 
centre (Fig. 305, &c.)i the pistil ia 
of iwuL'orpelB, right aiidlfH:; alter- 
nate with th(?8c is a pair nf stnmciis 
on the side nest the axis, niateUed 
liy anollior pair ou the op|iuailc 
si<le of tlio pistil, the four longer 
and interior atamens j altenute 
with these, and lower in insertion, 
n single stumen on eouh side ; next, 
overhipping in teativation. 
vith tlie two single stanicns and the 
two pairs : lastly, four sepals, alt^muting with the four petals as 
a wliiile, the anterior and (wsterior overlapping the lateral ones 
In the bud. Now the median ('. f. tlic anterior ajid [wsterior) 
pairs of stamens occasionally have their coutigiioiis' tllaments 
conjoined, as iti Fig. 397. If this were at all constant, the 
infcn'ncsj wutild uinhfiibtL-dly be that the ca«e ia one of diorisis, 
and Ihnt the flower as t<» its easenlini oi^ans la dimenius. This 
is apparently Ihc best osfpluiiatioii to he given. It assumes that 
the chorisis In normally coinplote in the andrrednm of Cruciferie, 
inat«ad of incomplete, as in Fumariacca;.' And this view is 
conSnnei] by the fact that the tne<iian stamens are simiilc anil 

four petala, of eoraewhat ' 
which ewK-ntinlly nllernat« v 

' The liypotheiiii here mloiitiil, » to tlie indrociuiD, ii that of Sioinhi'll 
(1830), ond «( IClL-lilcr |ln Fliir*. 1«W1, 1872, and Bllithcn.l. li. 200), rcpUdng 
ihil of Kiintli, liW). Sm., i-niployrd in former cditloiw. The ri'Jcrlcil view 
mnkri (lie tliiwi-r 4-nivrni» up lo ihc jiltlil, and tlic ilanieiM ft!) of one eih*l«. 
nltenmllnii wlili the fuiir )i#l>l*, Iht median >t«ii>nif (u in our vii-w ) iloublti) 
by i-hnriiii. KrauM and Wretathko {titnl >■ »liavo by Kicblcr) wouli) 
have tlic floral drelr* a-rnvniua init 4-nuTuuJ by turn*; the calyi of two 
li-ini-riiu* dtclei (which It plainly ia|; the L'uruUa of iinc 4-mi-ruu> iriri'te 

fli.1, ^ 


r, Willi |K»U|mi nr 



singlo in Sonc))iora and many fipccics of Lcpttlintn, in which the 
lateral or Bhort stamens are at Ihe same time alwrtive. 

37!). It iH quite |x)!4sible that chorisis may be extended to the 
corolla of the cruciferous flower, and reduce tlio wliole to a 
syminctrioal 2-nierous plan, and to congniity in the i>erianth 
also with Fumariaccn;. The only obstacle ia in the petals form- 
ing ft whorl of four where all the rest is 2-meroiis, for the sepals 
are manifestly two decussating pairs. Now the ine<1ian petals 
of Ilypecouin are deeply 3-lol>e(l. An almrtioD of their middle 
lobe woulil leave them almost two-parted : a little more would 
BC])arate them ; then they would imitate the four cruciferous ]>etals 
as in the diof^ram, Fifi:. 39o. Applying this view to Cruciferie, 
the blossom in the two onlers would accord in having a S-merons 
three- whoi'lc«l perianth, the first and third whorls meilian ; ' as 
also in the dimerous andnrciuin. the first whorl of which is 
lateral. The dilferenee is that in Fumariaccie the two members 
of the first whorl of stamens aiigmeut by chorisis into three, and 
tlie seccind is wanting, or is present only in Ilypecoum ; while in 
Cnicifenf the first whorl is simple (of the two short stamens) , and 
the second is doubled. In Funiariaccn! only the first whorl of the 
perianth ccnints as calyx, and 
the corolla is of two whorl.'j ; 
in Cnieifera', the first ! 
second whorls are (.■alyx, the (( 
inner sepals answering ti> the ^ 
outer i*etals of Fumariafreie. 

.IKU. Chorisis along with 
anteposition of utanicns is well =" =" 

seen in Tilia or Linden, at loast in the American species. In 
the?te the inilcliuitcly numerous Klauicns are in Ave clusters, one 
bfllm' each in-tal (Fig. :V.w, 31111). and there is o ijetal-like body 

Hllcnmlirg Willi llii-CBlyx-tiicnilH'ra ns ii wlmli-; Ihe short Mampni folUiwinp 
lU > S^iK'niuB i-irfb; th<'ii tlic lmi(t clntiicns nn a 4-nK'niUB i-irclc; lastly tlic 
tmi-rniK pymreimn. (i. llm.'l.iw (in Trims, Linn. Soc. ht. 3. i. 105) would 
luTv the fliiwvr 4-tiiiti>iih ti.v the oiipproHiDn iif tlir fifth mciiiliers uf n 
G-nuTnui< tjitv, flii>l a fiirtluT iiu|i|in'«iii>ii of linlf of tlic ri'miiiiiiiitt exterior 
■t«iiini-t-in.-l>-. &!'. FiiiHlly, IlKn' l> the inii.h I>etti-r-iniiintaiiii'<l viev thut 
thf eriieifcroiis tlowiT i* 2'iin>n>uii liiruu);li(iiil. nn t-xplaineil in the folluwlng 
paninii|>h. :!il>. 

I Thl» view wa* taken i>y Steiiiln-ii. lit Ann. .'ki. Snt. nor. 2. iT.T (IMO). 
Mill if eaw-nlialt.v ntiriKluei-il liy > Itikiiiiiin l>olnniiiI, Mvsviiajelf, in Itull. 
Soc. Iiup. Sat. Sl.isf. l»:± 

•anm <.t Ibe flnm uf Tltln Amerli' 
Ivlaihol atamcn-cliutcr Willi lli i-el 

in American Lin- 


in cacU duster witJi which the Btamens cohere. The explanatm 
by diorisis is Ihot each cluster, petal-like ImkIv iiicluiieil, is a 
iiiiitli|itioiitioD of one sUinicn. The djagram (Fig- '-iW) nctii- 
mtclji' flhoiTs thut most of tlie stamena originate fVuin the outer 
side of the base of the petal-like portion : this is most naturally 
explained by me<lian chorisis. The au|>erposition of Ihe chtsters 
to the petals will take the same explanation as that of Khauums, 
Vitis, &e. (Fig, 363.) That the andrteciiim ia here composed 
of the inner circle merely is partly confirmed by the alternation 
of the carpels with the clusters. According to Diicharlre,' the 
development of the andrtEcium in a Mallow inilicates a similar 
stnicliire ; for the whole nnit«d mass originates IVom Gve protu- 
berances, one before each forming petal and connected with it, 
this by collateral t^horisls forming a cluster of Siemens, and the 
five cinstcrs coalescing as they develop into a tube of fiiamcnta, 
each as in Fig. 485. Now Hibiscus and its near relatives have 
a naketl tip to the &tamen-tu1>e, ending usually in five teeth; 
and Sidaicen, as is most strikingly shown in the Caltfomian 
S. diploscjniha. has two scries of stamens, the outer (answering 
to those of Malva and its relatives) in five membranaceous pha- 
langi-s, eupcrposi-d to the petals ; the rather nunierous inner 
eeriee, more or less in phalanges, surmounts an interior Glamcnt^ 
tube. Whence it is inferred 
that these, and the five teeth 
terminating the column in 
tlibiscns, represent the in- 
ner stamineal cirtlu which is 
wanting in Malva, as it is in 

381. Tlie case of Pamas- 
sin would be explained as 
anali^us to that of Tilia, 
bill with the stamen -clusters before the i>etal8 wholly sterile, 
and of fewer divisions, while an inner circle of five stamens 

' Compira RviiiJua, IMi, &, Ann. 8ci. Kiit mr 3, iv. liS. Dacliirtre snil 
other* wliu ilraw fm-l>- uiHin meilian chorlili to explnlo «nicpnaltii>n, snd 
coniiclcr dial cungcnlul iininn pmTna ii, ink? ilic i<hiilitnget in ihnc euca. 
like the tingle lUnwna in Vitla, to lie >n iutwr p»n at tlie i«i>l iiwtf. Hut 
thii Yitfw appctra lo hate liail It* rUy, 

* any. Gi>n. llluiir. il. 41, GT. T&-82. Tlic petition at Ihe <!inwl< iNfore 
Hie pcUli in ParimU mi.l Malvariwiu bringi llic former into ^yninictrit^Dl 
■llcmailon arlih *ud> an inner tiamm^ircle ; Ijui it it nut m In Illbiaciu, 
wlili^li hki Ihe carjiti* Iwfnrc Ihu ivpali. 

FID. 4<W, A palal uf Pitniiu4* ■'■rnllnlina. villi ■ triiiU Uamlnodium baton It. 
yia. 4UI. IHjicriuii .If itii tlowai vt PatnaHla CvulluUiu. 


alternate with the petals forms the effective androecium. For 
the scale-like body before each petal, and even slightl}' 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.^ Wherefore, if 
these stamen-like bodies really represent a cii-cle 
of the androecium, it must be the inner one ; and 
that is the more probable view. 

382. Multiplication b3'chori8is in the gj'noecium 
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 y f 
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 

§ 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 man}* 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 properl}' outgrowths of the epidermis onl}', 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. trichnmata)^ that is 
structures of which hairs are the t^'pe. They ma}' occur upon 
the surface of any organ whatever. Their morphology is the 
moq^hology of cells rather than of organs. Tbey will therefore 
be most conveniently illustrated under Vegetable Anatomy as 

1 Eichler in Fl. Brasil., Sauyagesiace, & Bltithend. ii. 424 ; I>nide in Lin- 
luea, xxxix. 2.30. Eichler refers to this as a confirmation of Celakowsky's 
explanation of obdiplostemony by posterior displacement. (3G5.) 

FIO. 402. Pistil of Droeera flllfonnis with tricarpellary ovary (transrersely diyided), 
and six styles, i. e. three, and each two-parted. 




1 the Glossary' as respects tor- 

respects tlicir structure, and i 

384. But into some bristles, such as those of Droacra, n boV 
Jac^nt strntiiin of tissue enters, including oue or moi'c ducts or 
even some woody tissue. Prickles are of this class ; and from 
the most slendei-. which pnss into bristles, tliere are all grada- 
tions of stoutness and induration. Much outgrowths maj- even be 
fonned ui most regular order, as the prickles on the calyx-tubo 
of Agrimonia and scales on the aeorn-eup of Gabs, and yet have 
no mor[)hologicnl imiwrtanee. On the other band, true represen- 
tatives of leaf or stem may, by abortion and depauperation, Iw 
redueoil to the structure as well as the npi>earnnce of tricliomes. 
Examples of this are familiar in the papijus (answering to limb 
of the calyx) of many t'oiupoaiUe, and in the bristles which 
answer to i>erianlh in many Cj-peraeete. The scarioiis stipules 
of I'aronyciiia and of Potamogeton, the ligule of Grasses, and 
even the corolla in Plantago, are eqiially reduced to mere cellular 
tissue. So tliat the structural dilTerence between trichomes ant] 
ont/frotiri/ii ' is not at all al>solute, and the morphological distinc- 
tion most rest upon other ground than anatomical structure, 

385. Among the corolliiie outgrowths most akin to chorisis is 
the Crtiu-ii {Corona) at Silenc and allied Caryo|ihylIacca', at tlie 

jimetioii of the claw 
with the blade of 
the petals (Fig. 408), 
the analc^y and 
probable homology 
of which to Iho ligule 
of Urasses (Fig. 
150)iaevideut; also 
the many-raywl fik- 
"" •" mcntons crown of 

Passion-flowers (Fig. 401) . which consists of two or more scries 
of such outgrowths. In Supindus and some other Sapindaceie, 
these lignlar outgrowths or intenial appendages are more like 
a doubling of the jietnl ; as also in Erythrosylum, wlierc they 

' ThU u Ilic br»l EngllBli n«mc (or Hie Emrn/mini of ihc Rcmiani, llw 
SiaUoUrma of Wkmiing. &c. For the- i1cvi-loptin.-iil Hnd iii«i;u(*tim ol lliii 
(nbJCL'l, IPC ffatminjt, in KjiibciiliiiTTi Viilen»k. MnlJpl- 1»7H, an-1 n Ur|tw 
tnvlisp on RsmlflmlSun ill Ph«fier>iB«m». Cn|H>iihit|KU. Itt72. Al«i, I'hl- 
wonn in But. i>it. 18T3; Ci-lakuwBk}' tn How, Wi; aud Elihlcr"s note on 
EeuryrHun in DiaihcDdlagrammf, ). 48. 

FIO. WS. IN-tal .,f SIlniB Pcnnnj'Wiililrk. wltb It* cm™. 



are often more complicated in Btructure. They are always on 
the inner face, aticl are commonly two-lobed or parted. 

386. Similar etamincal appendages are well known in Cnsctita 
(Dodder) , in Larrea (Fig. 405) and other Zygophyl- 
lacese, and less conspicuously in Gaura. 

387. To exteu<I to them the name of Ligule may 
not be amiss, whether they are regarded as mere 
outgrowths of floral leaves, without ftirther morpho- 
logical relations, or whether they be, at least somo- 
limcs, inteq)reted as the homolc^c of intrai>etiolar 
stipules, as their ordinarily two-cleft form, and their *" 
coincidence in Erythroxylum with an intrapetiolar two-cleft stipule 

g 9. FonMS OP THE Tonus or Receptacle. 

388. Toms is the more specific and proper name. Receptacle 
is the more usual. (303.) A normal reieptacle of the flower 
would be that of Fig. 316, the apes of the floner stalk somewhat 

rface main^ (ov- 
the seNcral mter- 

enlargod, roundish or depressed, and 
ered hj* the insertion of the Be\eral organs, 
nodes which it potentially contains bemg 
undeveloped. As the raemhers 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 la the 
former is mainly for the sup|>ort of both an- 
drtecium and gyna'ciuin ; in the latter, as in 
Mjosnrus, mainly for the gjocecium only. 
The fall of the matured carpels reveals it 
as a vcri- slender or bodkin-shaped pro- 
longed axis. Of broadenetl forms, the Sti-aw- 
Inrry. even in blossom, aflbrda a familiar 
example. (Fig. 40fi.) In the same order, 
Unlius odorntus shows a very broad and flat wr 

receptacle : in roses, it is so deeply concave as to become the 
reverse of the Btrawberrj- (Fig. 407). being um-8ha|)cd with a 
narrow mouth, u|x>n which the i>etala and stamens are liorne, 

T\a. tie. Stsnieii of Larro Mtxloui*, wllb a coiKplcuooii llgiilnle appenilogo *t 
FKi. MS. Keceptacle o[ » itnirberTy la longltudliit] MCtloB. MT. Swue oT ■ ro«. 

Zl2 THE FiaWEB. 

white the pUtila Hoe the wslla of lite cavity, tbe base or c 
of tbia ravitjT nnawi-ring to tlir n{>cx ot the rirawlirrrj'. 

S9fl. KoniFlitnr* inh'ni'Nlcit an; Irntrthenetl betv(N?n certain 
VKm\>en. In Schizait'Ira, the rec^ptsf Ic, tMrclj' oblong in blos- 
Mom, lcni(llKiiit greatly in fruiting, bo as to scatter the caqiela 
un n long fllirunn axin. 

•1U1. In many Gentians, in Stanleya and Waroa among 
f 'ru(-ir<rfip. nn'l in tnoHt nynvXv* at C'lvoiue, tbe interaoiJe oT tbe 
rc<v-ptiicle tietweeo stameoa 
and pistil is developed into 
a long Dtalk to Ihu latter. 
Ojnnndropeifl (Fig. 40a) is 
like ita near relative, Cle<Hne, 
i'xeept lliat tins xery long stalk 
' haM the lower part of the 
Htamens adnale to it : the in- 
ttTiKMle Ijctween the corolla 
anrl calyx is Irnoad and slightly 
elevated (or in Cleome, &c., 
narrower and longer) ; and 
so the several floral circles 
arc B* it were dpnwd apiirt by this unusual development of 
r(!4^'plariilnr inlcnxriles. In Silene (Fig. 408) and many other 
jtlnntA of the I'ink family, an inli^rnodf between the calj-x and 
(»rt;ll8 la |irolonKe<l int4> a stalk or iSlipn.' 

■ Uttr* I* Ott gc-nrnl name of ■ (talk fnmivd by the rpccpiaclc or lonia 
pan of It. iif \i]f a csrpvl. T» illitlngiiUli li* panlrulor naiuri' in any cate, 
till' riill'in iriu ti'niin irv tnurt' or let* rmpltiyiil : — 

Tnw tnirini. fiir ■ Mli'u olilcli lK.-tung> to ■ ■Implo pistil iticlf (where 
It In liiJiiiriliiitnut wllh n pcllnlc). ntid i« nn part of the rctrcpUctc. u in Copti* 
iir (i'.I.IiJ.n'ml. 

tliv'ii-iinuK, wUstv tli« (tlpo la an Intern oik of iveeplacio neu t>clo» lb* 
lU'iui'i'luin, »■ Iha poil^tatk In iiimi! Crat'lfcne, C'Icuroc, and (lynaiiilropali. 

(liix'ii'iiunK, wlicn it I'lcvali-t bulli ilamcm and plitll, ni it icvmingly 
duo* In tliu \nvter ■Tlpo n( OynandrnpaU, Fig. 40(1. 

AiitiiiriiciRK, wlirn tlio Ml|w U a ■li-vvlqieil InlvnuHli' lielwn'n Ih« 
calyx and cofilU. a< tn t)ic link family. Viu. 408. 

Otiiisakk I* a tiTm proiwriy applli-d lo a ilinrt and com para lively hrmtd 
poUtuii III n-iv)ilac|p on wtilcli Iho ttynm-'lnm ml*, a* In Rni- ami Orantce 
(Vig. 411), llnundilnnimo, tingp, Ac. Tlii* may cxlt'iiil up U'twei-n tlipcar- 
pnU and |inM liiiii, iir lli>- iippir part In-runic a 

('*RnipiiiiRii. a nanm prnperly ajiplli-d lo a porlloa of rKvptwIc which 
ii litnliintri'd bi'lwivn Ih? curpcl* u a ci-nlral axis, u in Gnrnnluni (Fig' 
4111 and many UmWIIlfum. Fltr- 4tl 

nn, MS. Sacllon nf ■ Hnaar of BUihib rminiijlTaiilt'a, ilwwlng tb« (lips or 
rill, wb rknrar «r OTDuilraHi, wUb Hani circlai Hpaiai«d « 



392. Instesd of fonuing a stalk, the elongation maj-be continued 

between the carpels in the form of a slender axis, as in Gcra- 

nium (Fig. 410, 411), and in tlie carpophore 

|fe jr of the fruit of Umbelliferffi, Fig. 412. In 

' Geranium, this prolongation of receptacle 

extends far above tbc ovaries as a beak, to which the styles arc 
adnate for most of their length. 

393. In Nelumbium (Fig. 413), the gjnophore, or portion of 
receptacle above the stamens, ia enlarged into a singular broadly 
top-shaped body, with a flat summit, in which the pistils (a dozen 
or more isolated carpels) arc 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 hypogt/noiu, as in 
Orange, Fig. 414. Here it is a 
kind of gj-nobase. When adher- 
ent to or lining the base of the 
calyx, it is perigynout, as 
Uhanmus (Fig. 413, 416) and Cherrj- (Fig. 337) : when tarrie*! 
by complete adnation up to the summit of the ovarj-, it is f/'i/fg- 
HOM, ns in Comus, in Umbelliferie, &c. Not rarely it dindea 
into loltes. as in Vitia (Fig. S'O, 380) , in Periwinkle and most 
A[MH-ynaceouB plants, and in Cniciferte. These are termed glandt 
of the disk, anrl indeed are commonly glandular or nectariferous. 

no, t 

nacnlaCam. 411. Thennl 
le kxl< or receptacle, to whtcb (Ley w< 

Kltb truU 

rrfliD tlie pruliinesllon at tti 
ernily *IUcbe<l. 

rm, V2. Mstore (hilt of 0«mnrrhli», tbc two mrpeln »pllttlng awmy It 
tbeflllfiirm [irnlongmliin of the rewpiMle, nt i«rpoplioi* 

Plli. 413. The Inp-jbapecl nxept^le of NelomWum. wlUi tbe pl«fl>. imi 

FIG. 4 

1 upper f,i 



It is not possible by any direct domonstratioii to distinguish Iie- 
tweon sucli ni-odiictions of the rcceptixflp, which are dassed M 
helongiug to the axis, and suppi-esscd or undeveloped pliyllous 
oi^ana, sui'h as stamens, n'hich glands of the disk may BOIIU^- 
times represent. 

3i)5. HfponUiInm. Inspection of Fig. 41.^, 416. and 337.and 
compoiison with Fig. 339, will sngfresl an explanation differ- 
ent &OU1 that wliieli is generally 
adopted. Instead of reganlhig the 
calyx as iK-ginning on a level with 
the base of the ovar.v. and the cup 
as lined, more or less tliickly, hy 
an expansion of the itvcptacle (the 
iwrigjnons disk), 
the calyx may be 
understood to begin 
where tlib and the 
ovary liecome free 
from each other. 
rt'cc|itJi('li', insi»'fld 
of convex or protu- 
lierant. is liere von- 
cavc, has grown up 
around the ovarj", 
which, however, ia f^ce ttoia the cup in the earher cittil llf^res, 
butlmmersed in it in Fig. 3.'i9 and Ihe like, A comparison with 
a rose-hip, an apple, and a ix?ar much strengthena this inlcittro- 
lation. which ia rather laigelj ailuptcd at this day. at least 
ihcoivtieally. It was (K-rhaps llist pmpused by Link, who intro* 
diii-cd the appropriate name of IIvi'anthim. A hvpaniliium or 
hiii'itnthlol n'ceplaclc is. as the name l>etokenB. a flowcr-asis or 
ivccpiaclo develo|»ed mainlj' under the calyx. Tlie name is a 
gooil one, in any ease; and snch atructui'cs ns those of Culycau- 
thuH (Fig. 417-119). a rose, a pear (the lower part of whidi ia 
o'idently an enlargement of peduncle), and of Cactus-flowers 
(Fig. 317) . although quite comiiatiblo with the theory of adnatloD, 
arc more simply explaini'il by it.' 

' But, wlipiher the cum tre wi-il cllalinguLalinMc ur nul, il t>? no nifHiw 
fullowD tliat the m-eplucle i>U.vb auohn pnii iii D[|inEinni.'cBuf ]H'ri)i>'n.v anil 
of Irfcriur or parlly iiifi-rbr ovarj". Bucli « vlfw \t attrniti'd by atarv AM- 
oultiii than the otlii-r. UtiUiu tlic m«li»tiun o( an InvUiblc rrcpptkcb nuit 

FIO. ilT. Flowering braocln^QtljeaBtliiu. tin. VsrtkBlKclluuofUioura-diapeil 
rH»|iUirlii, Ih» Imbrlmtol bnwti or •rttb mi lu lurflm vul vkj. 418, Mature 


Section IV. Certain Adaptations of the Flower to the 

Act op Fertilization. 

§ 1. In General. 

396. The introduction into morphological botan}' of the con- 
siderations now to be mentioned should have dated from the 
3ear 171)3, in wliich Christian Conrad Sprengel published his 
cunous 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 exc^ellent illustrations of it. * The interest in 
the doctnne now prevalent is witnessed b}' a copious special 
literature, beginning with the publication, in 18G2, of Darwin's 
book on the fertilization of Orchids by the aid of insects.^ 

be invoked wlienever 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 bes^nning at the top of a long and simple tube. And if 
three or more of the floral whorls may be congenital ly united, why not these 
also with the remaining one ? Van Tieghem, in his Anatomic 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. 

^ C. C. Sprengel, Das entdeckte Geheimniss der Natur im Bau und der 
Befruchtung der Blunien, Berlin, 1703. Even earlier, Kcelreuter ( Vorlaufige 
Nachricht, etc., 1701-1700) 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 fertiliswKl by Insects, and on the Gootl Effects of Inter- 
crossing. Tendon, 1802. Ed. 2, 1877. This last contains a list of the papers 
and books which boar upon the subject, published since 1802. 

Other lending works and papers on the subject are, exclusive of the 
other volumes an<l papers of Darwin, more or less referred to hereafter. 

Treviranus, Ueber Dichog^mie, &c., in Bot. Zeitung, xxi. 180J3. 

Hugo von Mohl, Einige Beobachtungcn iiber dimorphe Bliithen, Bot. 
Zeitung, xxi. 1803. 

Delpino, Pensieri suUa Biologia Vegetale, &c., 1807. Relazione suir 
Apparecchio della Fecondazione nelle Asdepiadie, &c., 1807. Ulteriore 
Osservazioni sulla Dichogamia. &c., 1808-09, 1870, and later papers. 

Axell, Oni anordningama for dc Fanerog^ma Vaxtemas Befruchtung, 
Stockholm, 1809. 

llildobrand, Die Geschlechter-Vertheilung bei den Pflanzen, 1807, and 
other pai>er8. 

Hermann Miiller, 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 I*rofe88or Kemer, which describes arrangements in blossoms for exclud- 
ing unwelcome guests, has not yet reached us. It introduces the new terms 
Autofjamif and Allofjamy, defined on the following page ; the latter compre- 


397. Thft nibject, hen eonetdend m a port of morpfaotogy, 
muat be fiiilj- tiraled, as regards acts and processes, uudt-r [Jiysi- 
(flifgy. K^on' IliJog in the flower b in relation to fi-rtilisalioi] 
and fniHiflcaliim, tlireclly or iudirectlj-. This section is con- 
cerned <ntli tiiose fldaplatioiM uf structure by means uf wliicli 
a^ntM cxteraal to the bloseom are broi^ht into service fur its 

3'J*4. Lirma-tiH ami bLs successurs taugbt that the adjiistracats 
In benuaphrodite flowers were siieb, on Lbe wbole, as to se<:ure tlie 
a]^jli(futi(>n of tbe tx'"^'> *'^ i^ stauiena to the stigma of its pistil 
or piiitiU. Tbc present view is. that Uiis is douMlesa strictly ■ 
secured in (vrtain llowera of a lumjcrate number of s{)ceies, liut 
never in ail the flowers oT any t^iicli si>e(.'iea; that in ordinary 
flowers, where it may communl,v take place, it is not universal ; 
that in the larger rimiiber of species tliere is somcUiiog or otlicr 
in the floral stmeturo wliich impedes or prevents it. Some 
fluwent are adapu-d for close fertilization : some for cross fertiU- 
zaUon ; Bome for either. Here two terms need definition, vix. : 

Clo4e /erlilUation or Sflf-fertUhalimi, or Aulo/fomg, the applicflr 
tlon auti action of a flower's pollen u|(on Its own pistil : 

Vrnti frriilU'iiiun, or AUoffami/. the action of the jullen of one 
flower on the piHiil of some other flower uf the same S|M>cie8. 
ThlB may be near, as wben between flowers Viomc in the same 
tluKler tir on the same plant : remote, when between flowers of 
illntiiict plants cif the same itunetliate parentage ; most remote, 
when Itetween ditTerenl mces of the some species. Any tUiDg 
bcyonil this is hybridization, or crossing of si>ecie9. 


SOU. The dbctriue now maintained appears to have t>eeD first 
pmiMiiinded by Sjirengul in the statement tbat *' Nature seems 
to have wished that no flower should be fertilized by its own 
pollen," — a pmjMMition which Is nut wholly tenable, for tliere 
are blrMWuniH HixH'ially lulnpted to self-fertilizatiou. Il was re- 
nfflrnii-d 111 onr day by Uarwin. in a similar adage, "Nature 
abhunt tHTiKtual self-fertilization," — a metaphorical espression 
to which no efli.'ctlve excreption bus l)een taken. And the infer- 
ence wim drawn by him. tbat some itniKirtnnt good In the species 
miwL reanlt fVum propagation through the union of distinct 
lndivl<luals. and especially of individuals whicU have been dis- 
tlnirt IVir several or many generations. 

hunillnii driiamgamn, fcrtiiliitlon \iy tMilk-n of oilier flnwcra of tlie Mnw 
lilmir, mill X'livjaMn, liy (Hillrii from s flower oii tiiDilicr iilant. 



400. Tlie 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 proof of the proposition is an 
induction from a very great number of particular observations. 
That intercrossing is beneficial is a rational inference from the 
aiTa}* of si)ecial adaptations for which no other sufficient reason 
ai)[)ears, 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 continned by direct experiment.^ 

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 AnemophUous afad 
Entomophilous ; literally wind-lovers and insect-lovers, but de- 
noting wind-fertilized and insect-fertilized, according to the 
agent by which pollen is transpoited.' 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 sui>erabundance, 
incoherency, dryness, and lightness of the pollen, rendering it 
\Qry transportable by wind and currents of air. The immense 
abundance of |)ollen, its lightness, and its free and far diffbsion 
through the air in Pines, Firs, Taxodium, and other Coniferse, 
are familiar. Their pollen fills the air of a forest during anthe- 
sis ; and the *' showers of sulphur," popularly so-called, the 
yellow i)owder which after a transient shower accumulates as 
a scum on the surface of water several or man}' miles from the 

1 Darwin, The Effects of Cross and Solf-FertiHzation in the VegeUble 
Kingdom, rx)n(Ion, 1»70. American Edition, New York, 1877. 

'^ OmithofJiitoHa^ i. e. bird-fertilized, flowers are to Ik* ranked with entomo- 
philous. Tlie large blossoms of Trumpet Creeper (Tecoma radicans) and 
of Trumpet Honeysuckle (Ix)nicera sempervirens), and others, are commonly 
yisited and probably fertilized by humming-birds as well as by moths ; and 
other birds are known to play a simiiar part in equatorial regions. 



nearest soui'ce, testiliea to tLosc particulai's. All amcnlaeeoDB 
trees (Willows esc«?ptcrf),IIonip, Hojia, &c., are wind-fi-'itilizcd ; 
and. among i)orfect flowers, tliose of most Grasses, Sedges, and 
Plantago. In the lattiT Janiilies i-HiiL'L'iolly, llie Kiitliers ore pro- 
triiilert or hiing out in llie air onlj- when just i-eady lo discharge 
their i)olIen, ami are at tliat moment suspended on suddenly 
lengthened capillary dixKtping filaments, fluttering in the gentlest 
breeze ; and tlie stigmas 01% either dissecUHl into plumi^s, as in 
most Grasses, or l)C9et with copious hairs on which iwUeu is 
caught. Olio physiological adaptation, very common in the fol- 
lovring class, is not unknown among hcruiniihi'odite wind-fertilis- 
nlile Dowers, where it is im|K>rtant for securing intercrossing, viz. 
Dichngamg. It is best seen in the common species of Plantogo 
or I'laiitain. and is deseiilHMl lielow. (■108.) 

404. Insect-rertilUillile or enlomopliUous flowers are correlated 
with showy coloration (including wliil^. which is most showj' nt 
dusk), odor, or secretion or net-tar, often by all three modes of 
attradion to inseets iKtnibim-d. [Some insects, moreover, visit 
flowers fur their jmUcn, a highly nutritious article, and onlina- 
rily ptwiiiced in such abnndance that much niay be sjwi'ed. 
The showinosa of corolla or other floral envelopes is an attractive 
adaptation to rertiJizalion, enabling blossoms to Iw discerned at 
a distance ; nor do we know that fragrance or other scent or 
tliat nect;ir suliserves any other uses to the flower than that of 
alluring insects. Adaptations in the pollen of sucli blossoms 
for tmnspr^rtatioii by insects are vailons. Commonly the grains 
are slightly moist or glutinous, or roughish, or studded with 
projections, or stnmg with threads (as in (Enothera), so aa not 
to be rciLdily disiterscd in the air, but to have some slight 
coherence as well as caimbilily of adhering to the head, limbs, 
or bodies of insects, especially to their rough surfuoce; and 
in two families (Orchidncciv. Asclepiadaccie) the i«>l!cn is com* 
bineil in masses and with siwcial adaptations for being tntna- 
])orted «» muue. (421.) With tliis the stigma is usually 
correlated, by itmghuess, moisture, or glutinosity." 

405. Adaptations of the flower itself in rclbrence to insect 
visitation arc wonderftilly various ; and most of these are found 
npou investigation to favor, or ollen to necessilale, iutercrosa- 
ing. In dia'cious flowers, this is nceossitat^d by the scpnrulion : 
tn monireious an<l polygamous flowers, of various kinds and 

' Tliu* nrnrly vrcry l.>n.'liiit gi-nus but unc hut u iicnlalrnily iHutinuut 
•lifCiiiH : in tbtM-xifpUutikl one, Cypriptiiium. ii in tnoiti and minutely rough- 
Fucd, in eorrdation with [he loowly grnuulor or jiultkocuus pulUm which it 
j« to racciTc. 


degrees of separation, pollen is very commonly borne from 
plant to plant ; in hermaphrotlitc flowera only are more sj)ecial 
arrangements neede<l to secure intercrossing or a certain measure 
of it, and in these such arrangements al)ound. 

406. Irregularity is one of the commonest modifications of the 
flower (32G, 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.^ 

407. Dichogamy, a term introduced by C. C. Sprcngel, who 
first noticed and described it, is one of the most usual and effect- 
ual (ratlier physiological than moq)hological) adaptations for the 
promotion of intercrossing between hennaphroilite flowers. It 
means that such intercrossing is brought to pass by a diflerence 
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 

Proterondrous (or Protandrous) ^ when the anthers mature and 
discharge their pollen before the stigma of that blossom is recep- 
tive of |x>llen ; 

Proteroffynous (or Protogf/nous) , when the stigmas are in 
receptive condition Ix^fore the anthei*s have matured their pollen. 

Si/nanthesis,^ the maturing of the two sexes simultaneously or 
nearly so, is however made to secure the same result through 
special arrangements. 

408. ProterogD'ny. The Plantains, such as Plantago major and 
P. lanceolata, are familiar instances of this in a wind-fertilized 
genus with Iiermaphixxlite flowers. The anthesis proceeds from 
base to ajiex of the spike in regular onler, and rather slowly. 
While the anthers are still in the unoi)ened corolla and on short 
filaments, the long and slender hairy stigma projects from the tip 
and is receiving |>ollen blown to it from neighboring plants or 

1 This did not escape the attention of Sprcngel in the last century, and 
along with it the fact that strictly terminal and also vertical flowers, whether 
ere<'t 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 access!- 
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 whid-fertilized. 

'^ S*/narmtf is the term pn)posed by A. W. Hennett, in Journal of Botany, 
viii. (1870), 31(5, with its opposite, Ihteracmy, iov prut era udry and jtroterogyny. 
The latter names, in their shorter form {jtrotandry and protogyny), appear to 
have originated with llildebrand, 18G7. 


spikes : a <Iay or two afterwards, the corolta opens, the filam^nta 
givatly lengthen, and the four autbers now |>endcnl fl-om them 
give their light jxiHen to the wiini ; hut the stigmas of tliat flower 
and of all below it od that spike arc withered or [uiHt ivceiving 
pollon. Among Grasses, Anthosanthum is in the same case. 
The arrangement is somewhat similar to the Plantain in Amor- 
pha. which is fertiliued hy inseets, the simple stigma projecting 
beyoud the corolla in liuil, ivliile the anthers are still immature 

and enclosed. Scrophtiluria is a good instance of protorogj-rty 
in flowers fertilized by bees. The flower is irregular (Fig. 
417-41!!). and is approached IVom the front, the spreading lower 
lobe being the landing place. Fig. 420 represents a fVeshly 
opened blossom ; and Fig. 431, a section of it. Only the style 
tipped with the stigma is in view, leaning over the laiidtng place ; 
tile still closed anthers are enseonced below. The nest day or 
ft little later all is as in Fig. 422. The style, now flabby, has 
flillen upon the fVont lobe, its stigma dr}' and no longer receptive : 
the now-o[)ening anthers arc brought upward and forward to the 
position which the stigma occiiijied before. A honey-bee, tokli^ 
neutar fVom tlie bottom of the eorolla, will be dusted with poUoa 
thim the later flower, and on passing to one in the earlier state 
will deposit some of it on its fVesh stigma. Self-fertilization 
here can hai'dly ever take place, and only through some disturb* 
sncc of the natural coiirac. 

40!). Proterandry. The process is tJie reverse, and is at- 
teudcil with much more extended movements in Clerodeudron 
Thompsoniie, a Verljenaeeous tropical AfVican climber now com- 
mon in consenalorics. The ada|>tations in this flower (which 
we indicated long ago) are exquisite. The crimson corolla 
and bright white calyx in combination arc very crinspicnoas. 
The long flliform fliaments anti style, upwardly enrolled in the 

Fill 420. (21. Eailf dikikiI lluwDr «f l^ioiiliuUrlk iiwlo**. ftml ■ lu)iglluiUii4l 



hud, straighten and project when the corolla opens : the stamcne 
remain straight, but the stjle proceeds to vuvve dowiiwaitl and 
backward, as in Fig. 423. The anthers are now discharging 
pollen : the stigmas are immature and closed. Fig. 421 repre- 
sents the flower on the second day, the anthers elfete, and the 
filaments reciir\'ed and rolled up spirally ; while the style has 

taken the position of the filaments, and the two stigmas now 
separated an<l receptive arc in the very position of the anthers the 
previous daj'. 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 flying from 
blossom to blossom, and plant to plant, must transport pollen 
fl-om the one to the stigma of the other. 

410. Protcrandrj' abounds among common flowers. It is 
conspicuous in Gentians and in nearly all that family. But. 
while in Gentians the short style is inmiovahlc and erect, in 
Sabhatia it is thrown strongly to one side, ont of the way of and 
far Iwlow 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 es[K)sc the stigmatic«urface of their inner 
face in the place which the anthers occupied. In Sabhatia 
angularis, Lester P. Ward ' obsenecl that the anthers of fVeshly 

' In Mechan'i Gardcnort' Monthly, September, 1ST8, 2T8. 
FIO. 423. FIOwnofCkrgdgDdraiiThoiPEaanlM.llnt d>ri <2t,MCODdd*]r. 



Opened blosfloras ate all thrown to one side almost as stron^y 
aa the styk is thrown in the ojiposile direction. One or our 
common Fircwceds, Epilohiiitn auguatifoliitm or E. spicatuin, as 
it is variously c-alled, wliicli is common all round iJic novlliciu 
hemisphere, is similar to Snhlmtia in behaiior. In the IVcsIilj- 
o|>cnctt flower, white the.unthci'S are in good condition and are 

giving their pollen to Iwes, the etill immatnre style is atrongly 
cnrv»l downward and linekward. as in Fig. 42.1. Two or three 
days later, wlion the (lolk'n is mostly shed, the style straightcnB, 
lengthens to its full ilinu-nsloim, and spreads its four etlgmas over 
the line of the axis of the blossom (Fig, 426). in the verj' 
[xtsition to lie ixilUnaled by a hoc coming fi'om an earlier flower. 
411. In the following instances of prolerandrj', tlic style is 
made th*' instrument of dislrihuting the polk'n which it is not 
itaelf to use. The 
\f authors of a Cam- 
panula discharge all 
ir pollen in the 
unoiiencd bud, and it 
is nearly all deposited 
on the style which 
they surroimd, the 
upjier part of wliicb 
is etolhcd with a coat 
of hairs for holding 
In the ojien llowcr, the stamens are fonnd to be empty «n<l withered, 
as in Fig. 428. These flowers are risiU-d by liees and other insects 
for the pollen. While this is going on, and while the pollen is 
fresh and pleiitifiil, no stigma is apparent. Later, the top of the 
style opens into three (in some sjiccies five) short and spreading 
branches, the inner faces of which are the sliginaa. Althongh 


HO close at hnml, little if any of the [wllen of that flower can 

reach the stigmaa. These actually got fei-tilized by pollen 

brought hj' bees, which come loadwl with it iVom 

other Rowers and other plants. Sj'niphyandra differs 

from a tnic Campanula chiefly in the continued 

cohesion of the five anthci-s into a tube around the 

style. (Fig. 429, 430.) The pollen is discharged on 

an<l held by the hairy Hp|>er [x>rtion of the style. 

f"Oon after, the corolla expands, the lower 

]>ail of the style lengthens, and carries 

the pollen- load e<i part out of and above 

the anther-tube, as in Fig. 430 ; lastly, the 

three connivcnt tips of the style diverge 

and csposc the stigmas to iwilen mainly 

brought by bees from other flowers. By a 

slight further modification in Lobelia and 

in Comix)sitie, pollen is pushed out of the 

antiier-tube by the tip of the style as it 

lengthens, or by the vcrj- liaek of the two 

stigmas, the faces of which, aO^rwartls 

es|>ose<l, are not to receive this, but other 

pollen, though it may at times receive some of its c 

arrangement in Com|H)sita' is here illustrated from LeptosjTie 

maritima (Fig. 431-435), a showy plant of Southern California, 

now not \crj rare in cultnation The large flowers around the 


I. SUmpim and pltUl of > Toang, ui'l 130, i 
. Hewl of anircrt of Leirtoarnc marlilnia, nt r 

i« Troai «n old Bower ol 



mnrgiu (ray-dowers, with ligulate corolla), one of wliicli is sepa- 
rately ebown in Fig. 43^, are pistillstc only: the enUrgeil and 
estended open part of the 
? corolla (Iiright yellow in 
color) serves Tor aUrac- 
tion. the circle of raya 
gives the ai}|>earance as 
of a single lai^c dower. 
The flowers of the disk 
or whole central part are 
hernia ]ih rot lit«, au<l with 
narrow tubnlnr corollas, 
from the orifice of which 
projects the greater part 
of the tube of live co- 
alcscent antliers. The pollen is early discharged into the interior 
of this tube. The style, with somewhat enlargeil and bmsh-tike 
tip. at first reaches only to the 
bottom of the anther-tnlio : it 
slowly lengthens, pushes the 
pollen before it out of the tnbe 
(Fig. 433) and into the way of 
insei'ts of various kind, which, 
travelling over the surface, con- 
vey it to older flowers of the same 


hcail and of other plants. The style, elongating yet more, raises 
some of the (wllen still higher (as in Fig. 434) ; and at length its 
two branches separate and diverge (Fig. 4.15) ■ ex^tosiug to other 
pollen the stigmalic receptive surface which until now was un- 

412. !n Paniassia. which has sesoite stigmas, their receptive 
surface is netnally not formed until the anthers become effete ; 

no.«3 A llgii1iii«rpni>tofltmvtofthoutD*, anrl ■ antnl hcnnai>hnKtIte Bower. 
41X rpiwrpiul iif tlifl Uticr, iiiiifb «ilar|oJ, t]i« tubs uf kiiUisn prujei'iliig fmm Iliv 
imralk, ui>l U» pnllMi pmJwUng fnm apex oT Ibc anllicr-lube. bdng |>u*li«>t aptiy til* 
ItnfUmliigfofUigiilj'I'IwnMCh. Ml, Tlil* ntjrl* mi« pmjivtinc. and mm« pnllxi (HU 
rcnlnt on It* rip 4311. Tip of Muag (ij-li (laon ■ilianced and miicnlnBi])! Uii tva 
bnuiEhoi nimuullni, allll cartj'li"' '■<>■ «a Iba aiai uf cacli arm gr branch, anil 

bjr Iba dlTcrgoan now gipoaliif *Diin boa. 



and, as the plants or stems are single-flowered, the}' are fhnction- 
ally dicrcious while structurall}' hermaphrodite. 

413. The adaptations for hermaphrodite intercrossing with 
sj^nanthesis (407) , t. e, where there is no essential difference of 
time in the maturing of anthers and stigma, are manifold. 
They may l>e classed into those without and those with dimor- 
phism of stamens and pistils, or, in other words, those with 
Homogonous and those with Htierogonmu flowers.^ 

414. The cases without dimorphism are the most various, 
certain faniillcs having special tyi^es ; and are of all degrees, 
fix)m those that require intercrossing to those that merely favor 
or permit it. For the present purpose, having only moq)hology 
in view, it sufHces to bring to view two or three cases or tj'pes of 

415. Particular Adaptations in hermaphrodite blossoms, not 
involving either dichogam}' or dimorphism. These are exceed- 
ingly various ; but they ma}' l)e distinguished into two general 
kinds, namely : 1 , where loose and 
powdery pollen is transix)rtcd from 
blossom to blossom in separate grains, 
and 2, where pollen-masses or the 
whole contents of anthers arc bodily 
so transiKirtod. 

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 
to the stigma — 
were naturally 
supposed to be 

self-fertilizing; and so they sometimes are, yet with marked 
adaptations for intercrossing. None are less so than those of 

1 Terms proposed in Amer. Jour. Sci. »er. 3, xiii. 82, and in Amer. 

Naturalist, January, 1877. Dimorphism in flowers may affect the perianth 

only, and not the yov^ or essential organs; or there may be two kinds of 

flowers as respects these aUo, but with no rc»ciprocal relations, as in chisio- 

gamwis dimorphism (5.14) : or of two kinds essentially alike except in stamens 

and pisttil, and these reciprocally adapted to each other, which is htninjonoua 

dimorphism, or, when of three kinds, trimorphism. 

FUi. 436. Flower of Wistaria Sinensis natural size. 437. Same cnlnrpc*!. with 
sUndard. wliip*. an<l half the keel remove<l. 438. Same with the keel <leprt?N«e<l, as It 
Is when a Ue HliKhtH on this Its usual landing place, the cluster of anthere and Mtigma 
thus brought up against the iKse's alHlonien. 438. Style and stigma, with part of the 
orary, more magniHed, a fHnge of flne bristles around the stigma. 




IViatariii (Fig. 430-430), in which the light fringe of stiff haim 
ni-onm) the stigma (shovrii in Fig, 4311) would nut pa'veiit |Mi)len 
of siin-ouiiding anthers fi-oni fulhiig upon it. Yi»t when a 

ahghta upon Ihc keel, 
wit!) head townrd the 
1 base of the flower, 
I prolTOseis is inserted 
fur net-tar between the 
fool of the standard 
ml the keel, the latter 
1 is (li'pi'esBCcl liy 
■ eiglit, sfi Ibal the al>- 
(jinen of the insect i 
raiiiiht against the ten 
anthers fliid the stig- 
ma, becoming thereby 
smeared wilh pollen 
some of which when 
*" other Moi^soms arc via- 

itoil cannot fall to he applied to their stigmas. The very similar 
flower of Locnst (Hobinia), like that of the I'ea, adds an adapta- 
tioninfaroi-orintercrossing. Tliestyle forsome length below t ho 
stigma is covered with a short \xtatl of hairs, as is 
sven in Fig. 442. The anthers ojien early and dis- 
charge their i>ollen, which mainly lodges on tliia 
beard (Fig. 443). in a manner which may tbua for 
I>e Ukene<l to the case of Campanula. (411.) The 
wings and the keel arc yoked logctlier, and : 
together depressed by the weight of an alighting 
bee. Tliis does not bring out the anthers aa in 
Wistaria. These remain, early eflfete, within the sac, while the 
stigma and the poIlen-Iadcn {Mirt of the style (Fig. 441) are 
projected against the bee's abdomen, which, by the oblique 
mo4-ement, is first touched by the stigma and nest bnished ovei 
with pollen by the style below. So tliat, in (isiting a sucoession 
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 
(lower imineiliately gives to the same spot some of its jmIIch. to 
be tmnsferrcd to the next flower's stigma, and so on. 
417. Two special modifleations of the papilionaceous tj-pe 

>U|.l>lit. lha>lui<Ur<I >n.l olnK- n mnrixl. 411, ! 
«. raufllnf Uigillgmm mkI imllcn-lkilfli Up or Hi" 
m of wens In Ihe liuil. luting nns kml-prUtl, In 
. 44). Stjrie ind (tlgiiia U ■ Inter ihtW, Uw bnrd 


need particular mention. One of them, the Bean-blossom, i» 
well known to botanists ; the other not bo. The i>etiilianty iu 
the common liean, Phascolua vulgaris, antt its neai-est I'eiatives, 
1b that the keel, cnL'lusing the stamens and pistil, is prolonged 

into a narrow snout which is sptrallj' coiled (as in Fig. 444-446) : 
that the stigma is oblique on the tip of the style, and the beard 
on the Btyle is mainly on the same side that the stigma is : the 
wing-petnis stand forward and turn downward, forming a con- 
venient lamling place for becB. 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 
IVom first to last is concealed in the coile<l keel. Press down the 
wing-jx-tals. and first the stigma and then the pollen-laden tip 
of the style projects from the orifice : remove the pressure, and 
they withdraw within. 'When this pressure is made by a bee, 
resting on the wing-petals while searching for nectar vk-ithin the 
baae of the blossom between the keel and the standniil. tJie same 
movement occurs : the stigma first, and then the pollen on the 
style, strikes against a certain portion of the fVont or side of 
the Iwc'b body, and the repetition of this O|>eration causes the 
fertilization of each blossom by other than its own pollen. A 
slighter pressure or lighter movement of the wing-petals BufHees 

rin. 441. Rower or flitrclen Bon, Phacrolui vnlgnrta. 44S Suns with wtnR- 
prtsli prowl liiiwn ami U[i of nlTle prelecting fToTD Ihs niiflreorths kni. 44S. Some 
u 444 «ir*rR«l. mill «IHn<l>nl tnl wli<|^ reninTe<1. 4)7. Upper part of keel. In Ibo 
onmllilnn or 44.1. iml*npy|. iliowlng [.Ulnl; ihe priJecHiig Myla. 44i* Ssciliin of the 
keel, rn1ar«Hl. Khnwliig Ihe Hy]e oIlLIn bernre the anlhen open : lUtneni fbr Hke nt 
Hi«nH« nM ilelloculed. 440. PUtll deUcbed troni hi <Mm flowBr; Ihe broili loadad 


to jostle Bome of tlie potlen down upon ita own stigma, so that 1 

eolf- fertilization is not uacominon. 

41S. Apios tulxTOBa, a near reintive of Pliaaeoltis, exhibits n 

diltLTcnt and equally curious moclifleation of tiit! same parts. 
The wing-{>etals for landing [iWu 
are similar; the etandurd is pro- 
[xirtionally large, firm in texture, 
and sliell-sbapcd or concave, with | 
a small boss at the tip as seen tfom i 
behind, or a slialluw sac as seen 
from the front : the keel in narrow 

and sickle-shaped ; it arches across the fionl of the flower, and 

the blunt a{M<x rests iu the notch or shallow sac of the lip of the 

stnndanl. (Fig. 450. 452, 453.) So it remains if untondied until 
the bkissiim withers : no self-fertilization has ever been observed, 
and none ordinarily occurs. The anthers 
are assembled close aronnd the stigma, 
but a little short of it (Fig. 452) ; Uie 
jHillen is not early nor cojjiously shod in 
the enclosure : the small terminal stigniA 
is at first covered with a pulpy secretion, 
wliicli at length collects iutu & sofl ring 
, around ita base over or throu(;h which no 
' |iu1len jiasses. But when the keel is liber- 
attil by hfllngfrom nnderneulh, it eurvra 
prompt!}' into the shape shown in Hg. 

FIO *M Flower of A|4"» lubtma 
kwl dlslnil^ rrum the ntnlnlnf nntrh, 
trnileil ami Utrnal fnmni, fullovnl !•; tha antlien 

FIO VS. Eiiliii«BjTertlci»li*'llonnrflnw«»-l'mlufAii|»"'»w'™»- 
MUi iMir the Maiubfl ™i »"nr. to Phnw Iho Miint •pM at Oxb Ii«I : 
noMlu «4. lM«mni«filnw»r,*llhliiilrnftl»rt«n.t«rdcol»*«)',l"«h 
pliuK irhni tl» »|«x nf 111" U*t\ b llbofuWil. TlK flguiM (il*' lbo« fl 
|.n>wnt 1. kihI the lltM amnont uf ttM>dk|iUlbtwo(Apl». «u» publl>lia>l 
lo«n Asrlriilliitld l« ISTH. 

Iu till AnitT- 



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 hone3*-bees. In searching for nectar at the base of the flower, 
they probabl}' push forward into the space under the arching 
keel, and by slightly elevating dislodge its apex ; when first the 
stigma and then the anthers arc brought against some ()ortion 
of the insect's body, and against the same portion in succeeding 
blossoms, thus effecting cross-fertilization. This rationall}' ex- 
plains a i-emarkablc adaptation, which seems to be not otherwise 

419. Special Adaptations. Two of these, each ' peculiar to 
the genus, may hei*e be referred to. In Kalmia-blossoms (Fig. 

iU 4J6 457 

455-458), the anthers discharge the pollen through a small 
orifice at the &pex 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 1k>ss) of the corolla, in- a way analo- 
gous to that in which the keel of Apios is lodge<1 in 
the tip of the standaixl (418) : the expansion of the 
boi-der of the corolla in anthesis curves the fila- 
ments outwai-d and backward ; and when the Ik)wih1 «» 
stamens are lilK'Rited by rough jostling they fly up elastically, 
and the pollen is projecte<l from the two orifices. Some pollen 
may iK^ssibly be thrown u|K>n the single small stigma at the 
tip of the style, which riw»s much alH)ve the stamens. But the 
anthers are not dishxlgwl when undisturlKni, 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 

FIO. 45.'5 Vertical i«ectlon of a flower-bad of Kiilmla latifolia, nhowing the antlicni 
1o<Is<n{ in the Tx^cketJi of the corolla. 4M Ex|*aiuleJ flower, with bowed stamens. 
457. Vertical section of the same. 4SS. A stauien, cnlargetl. 

tiiutiT side of the abdomen fre(]iiei)tlj' touching the stignu, irMla { 
tin' prolwiaris ia senrching round the iHsttom of the flowor, Uberat- 
ing the stamens iu thu process, nhich one by one project Uieir 
|)i>lk-n u|»un the under side of the insect's body. In the iiassage 
IVotii dower to flower, ixillcn hi thus oonveyeil from the aiilbors 
of one to the stigma of another. 

420. Irifl luts tliree stamens, one before ench sepal or onter 
lulw uf tliu ix-riuuth, and behind each |>etal-Ul(e lulie of the etylc 

(Fig. 4it'J) : the stigma, ft 
efaelf-like plutv of each lobe, 
is just above the anther j 
but, as the anther favcs 
outward and the etigma is 
higher and faces inward, no 
pollen eaii find its way fktm 
tlie one to the other. But th« 
adaptation of parts is admir- 
abli' for couvc'jDiife by beea, 
wliit-h, siuuding uixm tlio 
ijidy landing (ilai-e, the re- 
curved Hfpnl. ihi-iiHt the head 
ilotvii lielow the anther, and 
in iidsiug it carry off imlteii, 
l<> lie allerwaitls ItKlgcd 
ii|K>]i tile tstigmas of other 
'" llciwei-s which they visit. 

421. Transportation of PoUlnla, or of all the |>ullen in a 
miuis. !■* elti'cUtl in most of Ilie species of two large onlera, 
not otherwise alliefl, the Asclepiadatn^tr aud the Orchidaceie. 
While in the Iris family the numlier of stamens is nxluced tVom 
six to threi', in nil the Orchis family, except Cj-priiKKliiun, Uio 
BtamenN are further reduced to a unfile one : hut the |K>llen ia 
|)eeidmrlv ■'<'oiiiimize<l. Ttmt of Anlhusa 1^4 in four livtse and 
soft |H-Iletj>, in an inveiteil cfts<pie-sha|M>d case. Iiingetl at the 
back, resting on a shelf, the lower face of which is glutinous 
atigma. oier the fnmt t^lge of which the casr|ue-HliH|)ed anther 
slif^htly projects ; and this anther is raised by tlir hcail of a Ik'c 
when escaping out of the gorge of the {lower. Tlic loose [tcllcts 
of [xillen are caught n)M-jii the liee's head, to tlie rough sur- 
face of whidi they are liahli' In adhcn- lightly and so to be inrried 
to Uic flower of another individual, there lefl n|K)n ita glutinous 

riti. Ua n«wer nf Iria |i>iirU«. vllh IWml pnrlhiii ■ml liitif nt nm tioUilnlJ mjlr- 
lalv ■■>■■ niani* cat ■«■)'. Tbi mcUun uf Uh aUtnu I* acn nlfcalM: Um rou|li 
Bppn (itrftcv ool; b nlituBilii. 


Stigma by the same upward movement which immediately after- 
ward raiscH the authcr-lid and carries away its pollcD, to be 
transferred to a third blossom, and bo on. 

4n. Hnt it is in Orchis ^ 

and in the commoner re- 
prescntatires orOrchis in 
North America (viz. Ha- 
benaria, &c.) that the 
most exquisite adapta- 
tions are fonriil, and the 
greatest econom}' se- 
ciirctl ; jiaralleled, how- ' 
ever, by most of the verj- 
numerous and variou8e|}i- 
phytic ami by various ter- 
restrial Orchids of wai-mer 
regions. A single illus- 
tration may here sulHcc ; 
and Darwin's volume on 
the Fertilization of Or- 
chids (ai»0, note), with 
its references to the 
copious literature of the 
subject, may be studie<l 
for f\i]l pailieutars and 
theirl)earings. Tlieflower 
is trimerous. ami the jieri- 
anth adnatetothe ovary, 
therefore ujiiuirently dc- 
veloiMxl II [ton its sum- 
mit, Tlie three external 
parts of the }x>rianth. 
whidi in Hal>enaria urbi- 
culata (Fig, iiiO) are 
much the broader, are the 
sepals: the three alternate 
and intenial, the i>ctals : " 

the base of the long and narrow petal which is tume<I downward 
is hollowc<l out and extended below into a long tiilw, closed at 
bottom, open at top (the spur or nectan), in whieh nectar is 


copioiiaiy BccrPtol and contained. The central part of i 
blosBuiii. bejoiicl tliP orifice of the nectary (shown scjMiratelr in 
Fig. 461), coiiaiats of one anther ami a stigma, fused U)!|fthor 
(the eliumtdrium) : tlie luarghial portiona. opening hy a U>ng 
ciiink, are the two cells of tho anther, approximate at their 
broader portion above, wiilely divergent below : most of the 
lower jHirt of the spatx between is excessively glutinous, and is 
the i^tigma. The grains of pollen are nnit<K^I by means of short 
Ihn'ails of very eJastic tissue into sniall masses, and these into 
larger, nud at length into pellets, hanng stalks of the same 
elastic tissue, by which tliey are 
all attached to a firmer central 
stalk, or *-ui«/ic/«. (Fig. 463-465.) 
To the lower end of this eandiele 
(directly to the end of it in our 
llak'naria; and Orchises gener- 
ally, in this instance to the inner 
side of the end, with & thick inter- 
mediate base inten'cning) . is aU 
UicUed a button-shniMHl ilisk, tha 
face of which is esi>o!teil, aud is 
on a line with the surface of the 
anther; so that these two disks 
look toward each other across the 
hruad intervening slignmtio space. 
aa seen in Fig. 461. The exposed 
face of the disk being covered with 
« tji a durable layer of vei-y viscid mat- 

ter, Uie body itself is sometimes tcnned agland, andimt improiwrly. 
The ^-iaciility is nearly of the same nature as that of the intcrveu- 
ing stiginn, of which tlic glands aiv generally 8U]i{)osed to be 
detached p«>rtionB. If so, then a portion of the stigma is cut off 
R-om the i-est and s|H.>cialtzed to the purjioseof eimveynnceoftha 
l>ollcn. When a finger's end or any sm.illcr body is tonched to 
these disks, they atliiere so fiinily that the atlached pallinia or 
|K)llcn- masses are clragged out of the cell and «inie<l away en- 
tire. Rome of tliese pollen -masses ha\'(' Ixwn found attached b)" 
tile disk to the eyes of a large moth. 'When a moth of the size 
of head and length of jtrolwscis of Spliynx dnipifernruin visits a 
spike of these flowers, and presses its head into the centre of the 


■numilliol |i-illiai-nMw> nr riatnTiUian iirMralktu. allli II> nhilk 

g nr Ih> naiKtiii* |»riL •«» -i |«llEn-|s>-k<i(ii. «l>h imm of llw 

Kilns Uwm MA. A pnnlon nimlilgJily macnlflod, wltli 

|Hil1>!U.gnii>i la foun i1<M4kIi«L 


flower so that its probisstis may reach and drain the bottom of 
the nectanterouB tube, a pollen mass will usuallj be affixed to 
each eiie on withdrawal, 
these will stand as in iig 
4C6. ^V ithin a nnniite the} 
will be Humed downward 
(Fig. 4C(i*), not by their 
weight, but b} a contraction 
in drying ot one side of the 
thick piece which connects 
the disk with the stallc 
When a moth in this con- 
dition passes from the last 
o[)cn flowir of one spike to 
ttiat of another plant, and 
thniets its pi'oboscis down a 
nectarj', the traiiHiiorted pol- 
len-masses will be brought 
in contact with the large 
glutinous stigma : on with- 
drawal, eitlicr some of the 
small iH'llcts of iHillcn will be 
left atlliercnt to the stigma, 
the connectiiigelustic threads 
giving way ; or else a whole 
)H)llen-mnss will be so left, 
its adtiesion to the glutinous *''•* 

stigma tieiug 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 poll en -in asses. In either ease, new pollen- masses are 
carried olf from fresh flowers and applied to tlie fertilization 
of otlicr blossoms on the same and eventually on those of differ- 
ent iiiilividiials. Cases like this, and liundre<lB more, all equally 
remai-kablti. st'ne to show how sedulous, sure, and economical are 
tiK adaptations ami processes of Nature for the intercrossing of 
hernial til roditc flowers. 

■i'2'i'. In Asclcpias, and in most Asclcpiadacen>. ttie whole 
c<»ntcnts of an antlicr-cell are consolidated into u solid mass, and 
this is ill 11 different way adapted for transjiortatiou to other 
flowers- TlnTc are ten such i><>llen- masses to eacli flower, their 

FIG. «S Front jwrt nf S|>livr 
thern m-hl.-ii1nri. nffl <inl In eMh «yp 
Met, tUowlnif cbe iilliin tnmrn di 

234 TIIK FMlWKIt. 

(':ni(li(-l(*H attachcfl in p:iir^ to \\\v (luiilil** ur !«••-« !• j i- • : 
shown ill V'i*i. U'1'1, 'I'lic ;;l:iniU :if-i- :illirri:i(i* h.!.. * .• . - 
ami NO (lit* two |M»ll('i|.iiiasM-<« mi*»)n-ii<Iii| U***.\\ •:»•.• j. . • 
loll;; to a«lja('('nt anther^. 'I'Im* a)i|i:ir:iiii'* i« iiHi^i 
aiitlirrs ainl traiis|N>rtnl liy vi^itiiitr iii^-«-i-». hIhii ■ • 
tongues lia|>|N'ii in Im' ilrawii tliroiiuli tin- « luiik 'i' " 

wliirli iiiiiiii-«liatrlv and linnlv cltiM*** iiihiii iIh- ii.< n. - - 

• • I 

H('n*«itivr action, wtiii-li lien' |ila\'^ lli«* part wliiiti :ii < '* 
|M*rtonn(Ml l»\ tin* adlicsiiMi ofu ;:lntiiioii'« <*uir:i<i-. 

rj't. IMniorphhin, /. «'. tin* caM* oftwiikiniU fi M"*- n <• • - 
licrniapliroditr. on the saiiir *«|NM-ir*«. i^ aiM»llii i :k<i:i;>* .* 
iiit('i'('n»s>in<^. Not all «liiiii»riiiri*«iii. ImufxtT. I'i'i iii*- ' •* -,■ 
diniorpjii^ni (l.'il) tlir intriil to M-ir-ti-rtiii/.r i-. i%:I<ii! I 
may al^i In* 4liiiioiplii*«in a«i to tin* |N-rian!li. h>'t | i'' 
atrcrtiii;! rcitilixatioii. ( Mic kind. lio\\«-\(-r. and tli« fn ::. 
is a s(N*(*ial adaptation to int«'n'ri>^**in;;. \i/.. : 

-IlM. llctrniiroiiouM llliiior|»hUni. i ll.'i. n<»ii-. ) In.- s 
applied t<' tlic caM' in uliii-li a «*)Hric*» pr«»*liiii-^ :%«•• k ■ > 
luTinaplii'tMiiti' tli>\vci->. oiTnpviii;;ditr«-ri*iil iiidi%idn:iU. iIm r' >^ 
c^M-ntiallv >iniilar r\(-i*pt in tin* anilrnTiiiin :iiiii 'j^iiot.i- 
tlicM> iTcipMH-ally ilitlrmit in IcnLrtli or liii;:lit. an>l :;.* i 
tioiis Hiii-li that, hv the aL'i'iirv of in**i'«-|o. Ihi' ■■•■ll* n !;• - 
•^taint-n-i of tht- tmr ^**i\ H'i'ipiiMall\ t'i rtili/.i"< tin- *! j'» l • • ■ 
other.' 'I'hi-« iiiiiiorphi*«iii ha^ Ih-iii di-tt'itiit ina'M.ut j.-:* j. - • 
iH-loii^ini: ti> t'lXii'tiM'n or tiltn-n natiiial i>idi-r'<. %i;<Iii\ •»• i"- -- 
thiiHiiih till' \f;^tla)»lc kiiiL'iliiiii : I>iit ilii-ri- an- t'ai !n<>ii • \..- 
aiiiiin;: thi' KiiMai im* than in aii\ ullur «>idt r. N«>ii]tt t. • « ^ 
th«- '*]N'rii'H of a LiciiU'^ all* ht-t«'iiii:i<ii"Uo. a*« in Ili'n»Ii>ii..k. •: 


' "1 1 I- .i« 'ii'ii u.i- ili-i ii»« rul \'\ Mr ll'irk .iii'! :ii* • -: ^a'- 
.1 4f n .' : .i!il I.'!«.iri| I'lill- "l I'hii.i.i. Ijil i:i .i- fij».r:.-i .;. !• ■ ■ . 
Ai .I'i N ir *». : Ai.ju-i l-7*». L". '•.'■. «li" I ;n« \.flii'l;: 

- I ' !- |.i . i;i. 'f i'»:inji iiii lit li;i» *•• • II l"iii: kn-'Wii !i. .1 (• « ; ;.*■■•• • . 
;»< P».'!-,i \.'- r jr iii.lit1-.r.i .iri>l M-n.-t'-rin hi l.-v.* .i' . •■-. 
I ! .'.I ■■: N- ■ ''i Ail.' r.' .1. i; '.> :'.'•. 1-1'. ■: I. • •• -a 1 :■■ '• 
li., . -'{.ii.-'i. !. .• !■ t.i'i'ij ll .1? till \ ;ir» ;it :ill ij»!i*i \i;.il l-i.! !' .i^ ' 
|. -Ill- .11 I i;!i -in. r- II' .■■■Ii\ I'lii lU '1 I.I .r iih :iT ill.' «.i- •!• (• s . ■. 
I ».ir« ;■■ :i!.l ! .i'!i k." ■ « •. n !■ - f' i;- r ' •■" T'-' ' "" I ■'?';- ■■'' I ' " 
f ,1 .; • ..| .1, • I . ^! . ■ ~ . i 1*1 : II I III. I .i:i>: .III ill' :r Ki k.iM*- >• x ' t I. a 

* I - . ■,!.' .■■ . ■; •, • . .: !;■■ il ii!" t!, I I Ill *».H . ;\ . *i . !•»*■- T* • ■ 

■ .. , ■ l-^: ,- ■! '.'.•. I- ij.'. • ■! 1 - \..!iiiiif I M'iMi •! ■ I ' • 1 • •'. ■ 
1 -'I.- ■ r" I ' M- •• • r •- ■ t ilii >.i"i. **i'« !• • "' Mr 1 »ir»ii ' I- *• ■ 
•■■.». :1 ..\. •- -.11,: ; / ' }•.'. 'i liM* * .".'.lilt ! •■ .».| J ■• ■ li 

1 • ,■ ■ . t •■ . :' // ■ ■■ I'-r !i I- k :: 1 'f I.; -...Ill Tit :.!T. r-'.. 

• ....;■ 1 1' ■ ' « " . ■ f ■ ':i" I'p! • \ 1 '. • • >. . • II :»• »i 'I .!• !'i. •' • 

« '. l' -r. M . I.'-!- "iij '■■■ 'I.' Ti iMit ! //■ ■ .« tiF //i.' tl.rr, r 

|>:..-;:i. a- iii« ii'.. -lit'! in .t l-.riiiir ti*>*.*. l\-'» 


Cinchona, sometimcB only a part of them, as in Primuhi and 
Linum. In Ilottonia, a l*rimiilaceous genus of two aixwies, the 
European one has hetcrogonous (limoqthisni * for cross-fertiliza- 
tion : the American one liaa honu^onoua showy flowers with 
only the general chance for intercrossing, and earlier flowers 
which are cleistogainuus fur self-fertilization. 

425. The natun^ of hetcrogonc <!in]or]>hisin may be well nnder- 
Mtix>d from a single example. The most familiar one is that of 
Ilouiitonia; but, in larger blossoms, Gelsemium is a fine illus- 
tration in the Southern Uuite<i States, and Mitchclla (Fig. 4G7) 
mostly ill the Northern, liaised from the seed, the individuals 

are almut equally ilivided between the two forms : namely, one 
form with long style and short or low-inserted stamens ; the 
other witli wliort style and long or high-inserted stamens. The 
stigmas in one rise to about the same height as the stamens in 
the other, )M>tli in tlie tall or exserted oi^aiis and in their low 
and included countcriiarts, as is shown in Fig. 468, answering 
to the left hand and Fig. 4C9 to tlie light hand flowers of Fig. 4G7. 
A bee or other insect with ])roljo8cia of about the length of the 
corolla-tidH:, visiting the blossoms of Mitchella, will brush the 
sume |iai-t of its iHxly against the high anthers of tlie long- 
stamened and (he high stigmas uf the long-styled forms; and 

' C. C. Sprergil, iw Darwin montions, liul notletd lliie, before 17M. He, 
" witli Iii* uaiihI Hnjtarity. aiiila lliat )ii.' diics not IkIicvc tlic e:(tal<'nce of 
tlie two foriiw lo Ih; *(;i.'iili-ntnl, lhuu|;h he cannot explain tlidr purjiusc." 
Darwin, Forma of Kluwi-rn, 51. 

Some lietcTOfionuiu I'rimulat aro «ai<1to proiluoc homoKonolu Taricliesin 
cultivation. In Priniula. anil in otlicr goncra, tlicrc are apecies which accm 
aa if of iini- Rcirt iiiilr, no recipnieal sort lieinK known, ai if one form had 
bci-umv Bclf-fcrtiic ami tlic other hail disappeared. 

riCi. 1ST. Partiiilge Brny, MllchslU repen*. In ths two fiiniu, itli. long-Manuoml 

and iliort-ity1c-[, aiul Btiort-atamenol and lonj^tylsil. 



the §nme [iflit of the proljcscia agnitiat the low anthers ofl] 
Bboi'l-stjimeiieti auti the low stigmas of the ahoit-stylwl funn. 

■l-iO. Moreover. Dai^ 
*in haa usccrtained by 
nki'oscoiiical exumina- 
ion that the pollen of 
f the tvto ditfcra in size or 
sliapG, and by experi- 
ment that it is less 
active upon its own 
stigma til an n^xin tho 
dttier ; indeed, that in 
many cases (as in some 
Bptfics of Linnm) it is 
qnite iiiaelive or imjio- 
tcnt not only npon its 
onn stigma but njMin its 
own-form stigma, whilfi 
it is prei>olent on the other, anrl this rcciprocallj- of tlie two 
forms.' Here, then, arc flowers structurally bcmiaphrudilo, but 
flinctionoll}' aa if di<»cioiis, securing ail the advantages of Uie 
latter, along with the economical advantage that l>oth sorts of 
individual and every blossom may bear seed. AVitli diwcism 
only sbont half the plants i.H>uld be fruitf\il. 

427. Hetero^nous TrlmorphLun. A threefold beterogooiem 
is known in i-crtnin siK-cios of a few genera ; and this ciimplica- 
tion may have certain mncci\able advnnt^iges over lUmorpliism. 
Wlicre scfdling dimorphous individuals are few and far between 
(those multiplying from root would all Ite alike), there would 
be an even chance that any two near each other were of the 
same Ibi-m and therefore sterile or imperfectly fertile. But if 
the organization were of thrte forma, any two of which inter- 
crosstnl with jwrfect ferlilily. Uio chances (as Darwin remarks) 
arc two to one tliat any two plants were of difTeivnt forms, and 
therefore by fertilizing each other completi'ly fruitful. 

428. The earliest known instaui* of three forma as to recii>- 
rocai relative length of stamens and pistil is that of Lj-thnim 

' ImiHiltiitc of own piillni, [-iliieT alwioliite or n'lmivi'. o 
ct^ain flnwrrs which arc not iKmorjihoiin, ut lii rnrvrlnli*. > 
PmtiflorB, Ac. On llie mntrary. Tiiaoy iliinorpliou" Briwpr* « 

I no Ic^s hi 

Mii|»n. VO, Lung^UniptiK] flower of 



Salicana. Thia was indicated by Vauther in 1841, more par- 
ticularly described by Wirtgen ill 1848, but was inteq>rctcd by 
Darwin, an<I tlie more recondite <Iifference9 brought to notice, in 
18C4,' " The three formB may be conveniently called, from the 
unequal length of their pistils, the long-styled, mid-styled, and 
ahort-styled. The stamens also are of unequal longtlis, and 
these may be called the longest, mid-length, and shortest. " 
The pollen of the different classes of stamens is of two soils as 
to color, and of thi^ee ns to size, the lai^st grains fVom the 
lai^st stamens. "Tlie pistil in each form differs from that in 
either of the other forms, 
and ill each there arc two 
sets of stamens, ditrerent 
in appearance and func- 
tion. Itut one sot of 
stamens in each form 
corresponds with a set 
in one of the other two 
forms. Alt<^ether, this 
one siMJcies includes three 
females or female organs, 
and three sets of male 
organs, all as distinct 
from one another as if 
they beloiigc<l to different 
S]H<cies : and. if smaller 
functional ilitfcrcnces are 
considered, there are five 
distinct sets of males. Two of the three hermaphrodites must 
coexist, and iHillen must be carried by insects retiprocallj- 
ftom one to the otlier. in onler that either of the two should bo 
fully fertile ; but. unless all throe forms coexist, two sets of 
stamens will ))e wasted, and the organization of the BtK'cies as 
a whole will Ik' incomplete. On the other hand, when nil three 
hermaphrodites coexist, and |>ollen is carried ft-om one to the 
other, the sctiemo is perfect : there is no waste of pollen and no 

' In an nrlirli- On tlie RoxurI Relations nf the Tlirw Formii of Ljtlirum 
Salicaria, in .Tr,iir. Linn. Sot-, viii. Idfl. Mm on the Cliami'HT and Ilybriil- 
like Nature of Ihi- (»tt?prinK of the IllrRititnati' Unimis iif Dimorphic aiiit 
Triniorphiir I'lanin. Ilji.l. x. 303, 1868. Reproiliited anil extemled in hit 
Tolunie enlilltd " Fomn of Flo»cr>," 1877. 

FKl. • 

■ T>lMnini nrtliB Ann 

irmi nf I.Tlhram Sallcarln. tn their 
■fl «n llic nnr dde. The .lottod 
:h pollen muat b« carrted to aach 


falBO co&daptation." The whole arrangement is displayed 
the aiinexeil (lingram (Fig, 470), and in tlie following aceunnt 
of tlie oix?ratioii.' " In a state of nature, thu flowers are inces- 
santly visitetl for their nectar by hive and other bees, various 
Diptera, and I^pidoptera. The nectar is secreted all round the 
base of the ovarium ; bnt a passage is formed along the nppLT 
and inner side of the fiowcr by tlie lateral deflection (not repre- 
sented in the diagram) of the basal ]x>rtion9 of the fllamcnts; 
so that insects invariably alight on the projecting stamens and 
pistil and insert their proboscides along the np|K!r and inner 
margin of the coi-oUa. Wc can now seo 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 bnished Iiy 
the lower hairy surfaces of the insects' bodies. The shortest 
stamens, which lie enclosed within the calyx of the long- and 
micl-stylert forms can botonehed only by the proboscis and narrow 
chin of a bee : hence they have their ends more upturned, and 
ther arc graduated in length, so as to fall into a narroiv file, 
sure to be rnkcd by tlic thin intruding proboscis. The anthers 
of the longer stamens stand laterally farther a[utrt and are more 
nearly on the same level, fur 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, they bend to that side 
where the stnicture of the flower allows the easiest access to it. 
as in Lj-thmm. . . . In each of the three forms, two sets of sta- 
mens correspond in length with the pistil in the other two forms. 
When bi'cs suck the flowers, the anthers of the longest stamens, 
bearing the green pollen, are nibbed 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 nibbed against the uuiler 
side of the thorax and between the front pair of logs. And, 
lastly, the anthers of the shortest Btamens and the sttgm.t of the 
sliort-styled form are nibbed against the (iroltoscis and chin : for 
the bees ui sucking the flowers insert only the fVoiit part of their 
beads into the flower. Ou catching bees, I otiserved niiidi green 
[wllen on the inner sides of the hind legs and on the abdomen, 
and much yellow |K>lieu on the under siilc of the thorax. There 
was also pollen on the chin, and, it may be presumcit. on the 
lirolioscis, but this waa dilllcult to observe. I hud, however. 

' All from Owwin, F«nin o( Flowpr., 13T-H7, Sc 


independent proof that pollen is carried on the proboscis ; for a 
small branch of a protecteil short-styled plant (which produced 
8ix)ntaneously only two capsules) was accidentall3' left during 
several da3*8 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 smaU 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 

429. Finally, a long series of experiments (requiring eighteen 
distinct kinds of union) proved that both kinds of pollen are 
nearly or quite imix)tent upon the stigma of the same flower, and 
that no ovary is fully fertilizable by other than a " legitimate 
union," t. 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 l)etween that of the long and that of the shortest stamens. 

430. Xesaea verticillata, a common Lythraceous plant of the 
Atlantic United States, is simihirly trimorphous, but has not 
yet been particularl}* investigated. Several South African and 
American species of Oxalis are equally trimorphous, and have 
been investigated b}- Darwin and Hildebrand,* with results 
quite as decisive as in Lythnim Salicaria. One genus of 
Monoc*otyle<lons has trimorphous blossoms, viz. Pontederia, of 
which the North American P. eordata is a good illustration.* 

431. All known flowers exhibiting reciprocal dimori>hism or 
trimoq)hism are entomophilous : no such wind-fertilized species 
is known. Few of them are irregular, and none verj' irregular : 
they do not occur, for instance, in Leguminosa*, Labiata;, 

J Monatsber. Akad. Berlin, 18(W ; 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 seetled freely." Forms 
of Flowers, I HO. 

2 Detected by W. H. Tx?ggett. 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 Ly thrum Salicaria, each flower having 
two sets of stamens, three in each set. 


ScTophulariaceEe, Orchi«laceiu, &c. Nature is not prodigal, i 
(Iwa not endow willi ueedkas adaptations flowurs wliicb are 
olliei'wiso provided for. 

S a. Adai 

i Closr Fertilieattox. 

4.32. Even where cross-fertilization in biaesual flowers is 
obviously arranged for^ it is apt to be tampered willi more or 
less of close-ftrtilization. The more exquisite the arrangements 
for the former are. the more completely is the plant dei^endent 
upon insect visitation. Failure to iuI^riTOSS is a remote and 
small evil c-omparcd witli failure to set seed at all. In order 
therefore that the plan of croaa-fertilization may not defeat even 
its own end, Uiruugh too absolute de|>endenee on precarious 
assistance, some opiwrtunity for seif-fei-tilization will usually be 
advantageous. Also there is a long array of insects visited 
flowers. es)>ecially polyandrous ones, in which close fertilization 
must lie much the commoner result, except where the pollen of 
another but wholly similar flower lias greater potency, 

433, Subsidiaiy- self-fertilization is secured in a great variety 
of ways. 111 Gcntiana Audrewsii, which ia proterandrons, and 
usually cross-fertilized by humble-bees entering bodily into the 
corolla, an exposed surface of pollen long remains fivsli u)>on 
the ring of anthers girding the base of the style : when the stigmas 
separate, they remain for some days simply divei^jent. 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 stjle. fila- 
ments, or corolla after anthesis commenoes. 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 
na Chirkweed. which Iiloasom through a long season, close- 
t'erl.ilize even in the bud in early spring, when insects are scarce, 
but are habitually intercrossed by insects in summer. Somewlial 
similarly, according to Hermann Mneller.' certain species, sucli 
as Euphrasia officinalis and Rhinanthus Crista-galli, haliitually 
produce two kinds of blossoms, one laiger ami more showy, 
usually affecting sunny localities, and with parts adapted to 
intercn^sing by iu»ecis : the other smaller or inconspicuous, and 
with anthers adjitstwl for giving })ollen to the aiyncent stigma 
without aid. There are gradations between these last arrange- 
ments, and the more special and remarkable one of dimorphism 


434. Cleisto^amy. Here the intention and the accomplishment 
of self-fertilization are unmistakable. This peculiar dimorphism 
consists in the production of very small or inconspicuous and 
closed flowers, necessarily self- fertilized and fully fertile, in 
addition to ordinary, conspicuous, and much less fertile, though 
perfect flowers. Two cases were known to Linnseus,^ 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 Malpighiacese 
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 MohP in 1863 ; but D. Mueller, of 
Upsala, who examined Viola canina, is said by Darwin to have 
given,' in 1857, '' the first full and satisfactory- account of any 
cleistogamic flower." The appropriate name of cleistogamous 
was given by Kuhn,* 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 
relationshij), though all but five are Dicotyledons. All but the 
Grasses^ and Juncus are entomophilous as to the ordinary 
flowei*s, and most of these such as have special arrangements for 
their intercrossing, either by dichogamy, heterogone dimorphism 
or trimorphism (in Oxalis) , or such special contrivances as those 
of Orchids. 

436. It has been said that the ordinaiy' flowers in such plants 
are sterile, and perhaps th€»y always are so except when cross- 
fertilized : in most cases they are habitually infertile or spar- 
ingly fertile. Probably they suflflce to secure in every few 
generations such benefit as a cross may give, while the principal 

1 Campanula (now Specularia) pcrfoliata and Ruellia clandestina, the 
latter a deistoj^amous state of R. tuberosa. Linnaeus did not make out the 
structure of tlie flowers, but supposed them to want the stamens. 

2 In Bot. Zeitung, xxi. 309. 
■ In Hot. Zeitung, xvi. 730. 

* Ibid. XXV. 65. The name (denoting" closed up" union or fertilization) has 
been written rl&istoifeno'is, which is not so proper. We prefer cleistogamous to 
deisioffamin (and so of similar terms), as best harmonizing with the Latin 
adjective form, both in fomi of termination and in euplioniously taking the 
accent upon the antepi'nult. 

* Amphicarpum (Milium amphicarpon, Pursh) is the earliest recognized 
cleistogamous Grass, except perhaps lA'crsia oryzoides. Some species of 
Sporol)olu8 are like the latter, and Mr. C. G. Pringle has recently detected 
such flowers concealed at the base of the sheaths in Danthonia. Amer. 
Jour. Si'i. January, 1878, 7L 



increase ia by cleistogamous Belf-feililization, which Ihus ofl^ts 
Uk incidental disadvantagt! of the fumier mode. 

437. In general, the cleiatogamous are like unto the ontituuy 
flowers arrested in development, some arrested in the almost 
fhll,v fonued bud. most at an curiier stage, and in the best 
marked eases willi considerable adaptive modification. Ia 
tUvne, "their petals arc rudimentary or quite abortetl; their 
staiiienH are often reduced in number, with anthers of very small 
Bize. containing few pollen -grains, which have remarkably thin 
trans])nrent coaUi, nn<l generally emit their tiibea while still 
encloHcd within the anther-cells ; and, lat-tly, the pistil is much 
reduced in size, with the stigma in some cases hardly at all 
devcloiKHl. Tlieae flowers do not secrete nectar or emit any 
odor : trom their Bmall size, as well as IVom the corolla bein^f 
rudimentary, they are singularly inconspicuous. Consequently, 
insects do not visit them ; nor, if they did, could they find an 
entrance- Suoli flowers are therefore invariably self- fertilized ; 
}'et they pnxliice an aliundance of seed. In several cases, the 
young capsules bury- Ihemaelvea beneath the ground, and tha 
sceils are there matureil. These flowers are developed before, 
or urtcr, or simultaneously with the perfect ones," ' In Grasses, 
however, as in some Dicotyledons, there is niucli less modifio»> 
tion and more transition. For when Leersiu half protrudes its 
panicle, in the usual way, the included half is fertile and the 
expanded portion sterile (or almost always so), although the 
flowers may open and exliiliit well-developed anthers, ovaries, 
and stigmas. But when similar panicles remain enclosed in the 
leaf-sheaths, they are mostly tVuitful throughout. 

4S8. Fully to apprehend tlie economy of cleistc^amj' in poUen- 
ssving alone, — and contrariwise to estimate tlie exi>ense of 
int«rerossing, — one should compare the small number of pollen- 
groins which so completely serve the purpose in a typical eleis- 
toganious flower (say 400 in Oxalis Acetosella. 260 in linpaticns, 
100 in some Violets) with the several thousands of all entomo- 
philous eross-fiTtllized flowers, rising to over tht^e and a half 
milhons in the flower of a I'eony. 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 pru<luction of 
odoi-ous material and uf Dcctnr. No 9|>ecies is alt^igetlicr tleis- 
togamous. Thus cleistogamy. with all its special advantage, 
testifies to the value of intercrossing. 


Section V. The Perianth,* ob Calyx and Corolla in 


439. The distribution of the floral leaves around the axis, 
which belongs to phyllotaxy, and their particular disposition in 
the bud (aestivation), have already been considered in Chap. IV. 
Sect. I., II. And most of the morphology of cal^^x and corolla 
has been outlined in the preceding sections of the present chai>- 
ter. AVhat remains chiefly relates to particulars of form and 
to terminology. 

440. Dnration. 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 calyx in Labiatse, in 
Phvsalis, and most Roses. 

Marcescent^ withering or drjing without falling away ; as the 
corolla of Heaths, Drosera, Ac. 

Deciduous, falling after anthesis and before fructification ; as 
the petals of Roses, the cal3*x and corolla of Columbine. 

Ephemeral or Fugacious, lasting for only a day ; as the petals 
of Popp}', Ilelianthemum, Purslane, and Spiderwort. In the 
two former, they are cast or early deciduous, the anthesis lasting 
but a daj' : 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 Cei'^us grandiflorus and other night- 
blooming Cactaceae. 

Caducous, falling when the blossom opens ; as the cal3'x of 
Popp3' and Baneberrj'. 

441. Nnmerlcal 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, 
b}' prefixing Greek numerals to the Greek name of leaves, ex- 
pressing them in Latin form, or transferring them to the Eng- 
lish. Thus 

DiphyUous, of two leaves (sepals or i)etals) ; TriphyUous, of 
three ; Tetraphyilous, of four ; Pentaphylhus, of five ; Bexaphyllous, 
of six, and so on. A tulip and a Tradescantia flower have a 
hexaphyllous [)erianth, but composecl of two circles, answering 
to calyx and corolla ; each Triphyllous.^ When the character 

1 Perianthium, alias Perigone or Perigonium. (296.) 

* As elsewhere explained, when numerical composition is indicated without 
reference to nature of parts, the terms dimerous^ trimerotu^ tetramerouSf pentO' 
imerouM, &c., may be used. (322.) 



of Uie organ, i. e, whether caljTt or cc)rolla is to he apccifie^i 
the word sepal or pelul is emplojetl in the oomhiuation ; as, 

Diifjioioiu, of two sepiiiB; TVifepaloiu, of Ihrec; Ttlrattpatou 
of four; Ptiitatefialom, of five (also written o-aepalous, ami SQ 
cordingly 2-sepHlous, 3-s«palous), and so ou : also, 

Dipetalout, Trlpelaloui, Tdrapelidom, Pmlaptta!ou$ (2-J 
petalons) , &c., when the corolla la coneemed. 

4-13. Monophylhut, Monoiepahut, and Monopetaloiu are 1 
proper terms for periantli (caljTt, corolla, &c.) L-omposed < 
einglf Ifaf. Likewise Poljfphglloutt PolyMtpalmUt and Polyf 
out for the case of n considerable but unspeeiBed niimher < 
members. Uiifortanately, in the Linnieaa and loug-prevale 
use. monopetalous was the term employ cd to designate a coroDi 
of one piece in tlie sense, or the fact, of a coalescence or groir'^ 
ing together of two, throe, five, or more petals into a cup or 
tube : and so of a calyx, of a whorl of bracts, Ac. And lioly- 
petaloiis. polysepaloiis. and po!.\"|)liyllous were the coiinterijarts 
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 Iw obsolete. In present 
descriptive botany, a i»l,\*t)hyllou8 calyx, or a poh-petaloiM 
corolla, or a 5-pctalous corolla, would be taken to mean tLat tl 
aepak or petals (as tbe case may be) were distinct or nncooKl 
bined. and a moiiopetalous corolla to be one with petals combinQf] 
by coalescence. (329.) 

-l-ia. Terms of linion or Separation. The pro|>er term for ^% 
corolla or a. calyx the leaves of which are more or less coalescenl 
into a cnp or tube is 

Gamopetahut for snch a corolla, Gamosepalniu for the calj:x fl 
these terms meaning united petals or sepals. Tlie older and miv-'J 
leading names Afonnpelaioui and Monotepahut. although current! 
up t*) a recent day, should be discontinued. Another term i 
not rarely uaeil in Germany, that of Sf/mpelalou*, for the gain^-J 
l>etulous (or formerly monopetnlous) corolla, — therefore ^~ 
tfpnloM for a similar calyx. It is perhaps a more apt t 
than gamopetaluus. and of the same etymological signification fl 
but the hitter is already well in use. 

Choripetaioia is, on the whole, the most fitting nai 
corolla the petals of which arc separate (us it literally expresse 
this), thot is, for what is still commonly called Pulyprtal'm 
already explained. (442.) It is adopted by Eichler. &c. C 
tepalaiur is the term applied to the calyx. Diaigpelatotii (en 
jiloyetl by Endlidier) bos the same meaning. Both this t«ri 
and choripctaloua carry the implication of separated, rathel 


than of typically separate, parts. Eltutheropttahui (literally 
free-petal led) has also been used, but is incoDvenieutly long. 

444. DegL'ee of coalescence is most correctly expressed by the 
phrases united (coiiuate, or coherent, or coalescent) at the bate^ 
to the midiile, or to tlie lummit, as the case may be. But it is 
more usually and tersely expressed in botanical description by 
employing terms of division, identical with those used iu describ- 
ing the lobing or toothing of leaves and all plane oi^aiis. 
(144-188.) That is, the caljx or corolla when gamophyllous 
is for description taken as a whole, and is said to be parted 
{3-parted, b-parted, &c.). when the sinuses extend almost to 
the base ; r/«/}, when about to the middle ; lobtd, a general term 
for any considerable sei)aration beyond toothing ; dtniate or 
toothed {3-loothed, b-loothed, &c.), when the union extends 
almost to the summit; entire, when the union is complete to 
the summit or bonier. 

445. Farts of Petals, Ae. The expanded portion of a ]>etal, 
hke that of a leaf, is the Lamina or Blade : any much contracted 
base is the L'Kctis or Claw. The latter is very short in a rose- 

petal, but long and conspicuous in a pink and all flowers of that 
trilw (_Fig. 471), in many CapjMirideie (Fig. 409) and Crucifene. 
A sepal is veri- rarely distinguishable into lamina and claw. 

44G. Parts of Gamophf lions Perianth. The coalescent portion 
of a corolla, calyx, or of a perianth comiKised of both (such as 
a Lily or Crocus-blossom) , so far as the sides are parallel or not 
too spreading, is its Tube : an expanded terminal portion, citlier 
divided or undivided, is the Limb or Border. The limb may 

PIG. IT1. Cnrnlln nf Sna[>*nrt. af Bts Hpanto long-daweil or uiguleulalt pctali, 

Fia. 4T3. Floirer nt Gllis caninoMMlii ; Iho puts answering to tbe cla<iO iit tbe 
petals nt llie lui flfnirr hers all nnltfd Intn a tabe. 

FIG.4n. F1nwcr<>rtbaCyi>r«is-VlM(Ipaiiui!aQniuunr11<):lheiKUlsaU(tle(arIlier 
inilcl Intn a flvp-lnlml ipmillng bonier, 

FIG. *T4 Fl'.itrrnrthoIpnnnpapoiTlnes; (heflTei-nmp^eiitpetiilFipprdvllrnnit 
Dio a trumiiet-Bliapei] tube, aud bejoul lutu an almiMt entire spremtlng border. 


be [tailed {tbat is, tbe compyiient parU not united) qaite { 

irlj-down to the tulje or base, as in Fig. 472, 47. 
as ill Fig. 473, 47( 

j-lolwd) ; 

r less a 
(with ] 


L inefel}' aoglesA 
or ixiiiita to reiii-caeiit the tips 
of the comi>oncni members, as in 
Fig. 474 ; or with even anil entire 
border, as iit eommon Muruiug- 
Gloiy, Fig. 482. 

447. Tlie line, or sometimes a maiiifest or (conspicuous iKjrtioii. 
Iwlweeu tbe limb anil tiilie (in the corolla alwajB a portion ubovv 
or at the insertion of the stamens, when these are borne by tbe 
corollaj is called llio Thkoat, in Latin Faux, pi. fuuce». This 
is mostly more ojien tlian tUc tube, yel loss expanded llian the 
limb ; but it otU'n presents insensible gradations fiom tbe one lu 
tbe other. 

448. Such appendages as the Coroha or Cbmwts {3ft5, shown 
in Fig. 103, 404, 471) usually belong to the throat of a gamo- 
petalous coroUa or periantli, as in Oleander, CoinJVey. llornigf. 
Narcissus, &c,, or to a coiTespouding position when tbe parts 
are not coaleseent. 

44!f. Forms of Corolla, Caljx, kc. As to terminology, Bome 
of tliese are sjieeial and are applicable to corolla only, as the 

Papilionaeeoui, the i>eculinr irregular corolla of the typia 
portion of Legiiminosie (388. Fig. 342-834). which has I 
already illustrated, and in which the t*etala, two pairs and i 
odd one, take ijarticular names. Also the 

Caryophyliaceoui, or Pinli-llower (Fig. 471), a regular coro 
of five long-claweil {iinguic^iliilf) petals, the claws enclosed infl 
tubular cjilys and the blades spreading ; and the 

Crticiftroiu, of four somewhat similar pctiils, the four abrapl 
spreailing blades in the form of a cross (cruciaU) , a.a in Fig. 394 

Jiotartovt, with roundish and widely spreading petals on V 
short or haiilly any claws, as in Rose and Apple-blossoms. 

Ziliaeeous, a 6-phyllous perianth of eampanulate or fbnrielfonn 
shaiw; the members either distinct, as In most common lilies and 
tulips, or gamophyllons, as in Lily of the Valley. All but the 
first and hist of these sorts are examples of regular and chori- 
])etAkHi8 perianth. 

OrehUfactoii» fiowers aw of a peculiar irregularity, eomlnning 
hotli catys and corolla : one member, tlie petal in fVont of the 

BoMte or «hMil4hai>ed ud llTii-partal c«roU> ot Uie BittenweM (Sd- 
Wli«]I-tLiip«J and STe-IOboJ coroUa of Uiu (Uiunina PoUito. 




Stamen and stigma, differs ttom the rest in shape and in being 
nectarircrous (as in Fig. 460) ; it is named the Labelllii. 

Gaieaia ia a term applied to a corolla the upper petal or part 
of wliick is arclied into the shape of a casque or helmet, called 
the Galea; as in Aconite (Fig. 357) and Lamium, Fig, 479. 
In the former the galea is of a single petal ; in the tatt«r, it 
consists of two, completely united. 

450. Gamophyllous forma with special names are chiefly the 
following. Illustrations are usually taken fl-om the corolla, but 
the forma and terms are not peculiar to it, excepting the firat, 
viz, the 

Liguhle or Slrap-$haped corolla (Fig. 288, ic.), which is 
nearly confined to Composite. Here a corolla, formed of three 
or fire petala, imitates a single petal, except at its very base, 
which is commonly tubular : the remainder is as though the tube 
had been split down CO the upper side and flattened out. The 
corolla of Lobelia, type of a family moat nearly related to Com- 
positie, illustrates this. (Fig. 488.) 

451. The names of the general forma arc mostly taken ftom 
some resemblance to common objects. All those In common 
use will be found in the Glosaarj- : a few leading onea arc here 
speciflwi. They may bo divided into the r^ular and the irregu- 
lar. Tlie principal irregular form with a apecial name ia the 

LnhiaU, or lipi>e<l, alao termed Bilabiate, as there are two lips, 
an upper and a lower (superior and inferior, or anterior and 
posterior, 290), although one of them is sometimes obscure or 
abortive. Thia bilabiate character in the corolla, and often in 
the calyx alao, per\adea several orders with gamopetaloua 
flowers, and gives name to one of them, the LabtatiB, to which 

via. in. CuBiannliite corona of rb« Hurebcll. rrnnpnnaU n'tntiillfnlin. tn. 
S^verfnrm (hypocnKeriDinrphnns) mmllB nf Plilnt iro. LaWnte IrinKentJcorcdC "^ 
Lamlnm; a ({.to Hair. tee. Pi-n»nale corolla oT Antinhlanm or Snapdraam. 
PcniHiate eorollm of UoarU, ipurred (nkante) at Uia bUB. 


the Sage and Mint belong. Sucb flowers are a-meroiis. 
Lave two merulwrs siwrially united to form one lip, and tlipee to 
the other. The odd seiMil being [wstvrior (or next the axis of 
inflgrescenre) , and consequently the (xld jjetal anterior, the (Td^Tt 
lias its lower U[> of two sepals ami its upi>er of three ; nhilo the 
corolla has its upi^er lip of two petals and ita loner of three. 
Uut in Leguminosic, where the cal}-x is sometimes bilabiate, and 
where the odd sejial is anterior (or toward the bract), this is 
reversed, and two sepals or lobes of the calj'x form the upper 
Up and three the lower. A bilabiate corolla is 

Riugent, that is gaping or ojien -mouthed, when tlie throat fs 
IVecly oi)en, as tn Lamium, Fig. 470 ; 

Pertanaie, or masked, when the thioat is closed, more or less, 
by a projection of the lower lip called the Palate, as in Aotir- 
rttinmii and Linaria, Fig. 480, 481. 

Ab'i. Of regular forms, there are the followlDg, b^inuing with 
that haWng least tube : 

Rvtate, or Wheel-tfiiiptd (Fig. 475, 47G) , widely 8|)reading from 
the vcrj' Imse, or frum a short aud inconspicuous lube. 

CralerifiH-m, or Saucer-iAaped, like rotate except that the broatl 
- limb is cupixrd by some upturning towaitl the margiu. 

Hgpoerateriftinit, or rather (not to mix Latin and Greek) 
HifpocraUrimorplioui, in English Sali-erform, when a rotate or 
8aucer-8ha|>ed limb is raisei! on a 
slender lube which does not mudi 
enlai^c upward : that is, where a long 
1 narrow tube abniptly expands 
into a Hat or fluttish limb, as in 
Fip. 478. In Fig. 472-474 are seen 
salvcrform corollas with somewhat 
more upwardly dilated {Irumptt- 
thaptd) tuljc. The salver or hi/pif 
cralcriu/n, which the name refers to, 
with a stem or handle beneath, is now 
to lie met with only in old pictures. 

Tubular, when strictly used, deuotea 
B gamophyllous perianth with Urab 
" inconspicuous in proportion to tlie 

tulic. as in Tnmi|iet Iloneysiiekle. or as Fig. 472-474 would he 
if the tinib were much diminishol or wanting. But it is some- 
times used in the sense of Imiing a cotispieuous tulw. 



Infundibidiform, or Funnel/ortn, such as the corolla of oommoD 
Moniiug-Glor)- (Fig. 482), denotes a tube gi-adually enlat^ng 
upward from a narrow base into an expanding border or limb. 

Campanulate, or Bell-shaped ( Fig. 477), denotes a tube of length 
not more than twice the breadth, moderately expanded almost 
from the base, the sides above little divergent. 

Section VI. Tue A]a>B<£CiCH, ob Staheks in pakticular. 

463. The whole Stamen. For the general character and some 
of the mo<liGcationB of the stamens, see the first (301) and por- 
tions of the succeeding sections of the present chapter. The 
terms |>eculiar to these organs, and of common use in botanical 
description, were nearly all coined by Linnieus, and employed as 
the names of chisses in his sexual system. (C72.) The sub- 
' stantive names of those classes which are characterized by the 
number of stamens, and which were designated by Greek nume- 
rals prefixcti to andria (the Greek word for man being usc<l 
metnpliorically for stamen), are put into adjective form, as 
follows : 

Munandrout, for a flower with a solitary' stamen ; Diandrout, 
tot a flower with two stamens ; Triajidrout, with throe ; T^ran- 
droua, with four ; Penlandroitt, with five ; Hexandrout, with six 
Jleptandrout, with seven; OctaR(A*ou«, with eight ; JSnneandrout^ 
with nine; Decandrout, with ten; Dodteandrout, with twelve 

Polyandrous, with a greater or indefinite numl)er, or Icosandrout 
(meaning twenty- stamenetl) when a polyandrous flower has the 
stamens inserted on the calyx, as iu the Cherry (Fig. 337), 
Pear. &c. 

Tin iK. TilKilflplinaaiUmeiialSuiil tjafa P««. IH. Monaddplioiu atuiieiia of 
■ I.oiilTW. twi. Mniuvlclphnuaauinena, Acrif MBllnv. 

Fio.4W. FicsiyDCtneiloiuiUmeiuDfiCompaiitA. 1E7. Xlis umB, l&ld open. 




out is a term ni)i)lie(l to an aii(IroH:iiiin of fou 
ineiiH ill two pnira, a longer and a shorter, ns in Fig. 

Telradi/namaui is similaily njiplit^d to that of six sUineus. two ol 
them shorter, intlie manner cliaraet«ri8tic of CruL-iferiB. ' 

A!tA, Tcrma vrliich ileuotv coalcBccoco of stamctui, nhctUer by 
their fl]um<<iita or their antliers, are 

Mimnddphotu, that is, in one brotherhood, by coalescence or \ 
the filaments into a tube, lis in the Mallow (Fig. 4So), LupitM | 
(Fig. \»\). Lobelia (Fig. «8), &c. 

Diadelp/iout. in two bnillierliuoils, by eoalesuence of the fila- I 
mentjt into two sets ; sometimes an equal number in each, rs in 
Fiimariaceie (Fig. 3'JO), sometimes nine in one set and one 
BG|iarate, as in the I'ea (Fig. 4M3) and most I'apiliouacea'- 

Triadelphom, with filamcnU united iu tlirce seta or clusters, u 
In Iljiwricnm. 

I'tntadtlphoM, in five sets, as in Linden, Fig. S98, 399. 
But in general, wlien the sets are several, without regard to the 
numlK-r t)ie stamens are said to bo Polj/ndelphout. 

Synffeneiiotu, when the stamens ore unltt'il by their anthcn 
into a tube or ring; as in tlui whole vast order of Composite 

(Fig. 48(1, where Iliey ore five in nnmber and the filaments dis- 
linet), in Cni-nrbiU (Fig. 489, 400, where they are three in 
uumlier and the fllaments jiarlly monodelpUoua) , and in l^liella 
(Fig. iSH. where tUoy arc also five and the long lilaments are 
mainly monatlclphous) . 

it liobolla uMlnalU, slUi tnbsnT rornlU dlrlrlol nn mifi dile; 
anlUilliifaiiiluhv^/iutvtirninnisiiU! <l of ■iiiluirx 

CiMBrMU (SiiUMlil. vrllb 11ni1> <^ «lyt bikI itiit»]U rut 

nntafgwL nii.l Uw npiMr p^rt mil ■w»t, in tln-t tb» onion. Tl» uillinn an di 
4>1. A ilrUrh^HMMnorilM Melon, wliLlnwtlTdnii-iiiiknUMr. 

Kig. tin. stittuiu Kn.l Xyla uf ■ Cyrv1|>"llum. unlMl Into oat bwly ot « 


455. Of terms relating to adnation of stamens, besides the 
general ones of hypogynousj perigynous^ epigynous (332), and 
epipetalotiSy or adnate with corolla, there is the special one of 

GynandrouSj having stamens borne uiK>n the pistil, as in 
OrehidaceiE. In Cypripedium, the filaments of two stamens, and 
an enlarged sterile stamen behind, are adnate to a style, while 
the two anthers are quite free (Fig. 492) ; in the prosier Orchis 
tribe (as in Fig. 4G0, 461), anther and stigma are consolidated 
into one mass, and there is no evident style. 

456. A complete stamen consists of Filament and Anther. 
The latter is the functionally essential part of the organ, and 
therefore is wanting onl}' in abortive or sterile stamens. (345, 
352, &c.) The filament, being onl}' a stalk or support, may be 
ver\' short or wholly wanting : then the anther is sessile j just as 
the blade of a leaf is said to be sessile when there is no petiole. 

457. The FilameDt, although usually slender and stalk-Uke, 
assumes a great variety of forms : it is sometimes dilated so as 
to resemble a petal, except b^-its bearing an anther ; as in the 
transition states between the true petals and stamens of N^-m- 
phiea, shown in Fig. 318. 

458. Such petaloid filaments would indicate that this part of 
the stamen answered to blade rather than to footstalk, while 
others would harmonize better with what seems at first sight to 
be the more natural view, that the filament is the homologue of 
the petiole, the anther of the blade of a leaf. Remembering 
that in large numbers of leaves there is no distinction into petiole 
and lamina or blade, such homologies should not be insisted on. 
The filament may be variously' ap{)endaged by outgrowths. Some 
of these api>endages are verj' conspicuous, such as the scale of 
Larrea (Fig. 405), which is on the inside, and the nectariferous 
hood of Asclepias on the outside ; or there may be a tooth on 
each margin, as in sixicies of Allium. 

459. The Anther, the essential organ of the stamen, contain- 
ing the iK)llen, suimounts the filament, when that is present. 
It normally consists of two cells or lobes, the word cell being 
here used in the sense of sac. But, as each sac is not rarelv 
divided into two ca^^ties (locelli), the best technical name for 
anther-sac is that of Theca. The two thecse, lobes, or cells are 
commonly connected b}' a more or less evident and sometimes 
con8i)icuous common base or junction, which is mostl}' a pro- 
longation of the filament, the Connecttvum, or in English 


460. For the discharge of the pollen, the cells of a uonnal 
anther open at the proper time by a line or chink, usually 


extending (Voin top to liotlom (Fig. 493), the ttUuri^ 
flr/iiicfiief. C^omnioulj- this line is lateral or inai^nal : not ' 
rurelyil faces forward or Iwdjward. In the vaut genus SoUuum, 

to wliich the Potato belongs, in most Ericaceous plants (Fig. 

458, 494), in Polygala, ami in tnaiiy other flowers, the anthor- 

uells i>i»n oniy by a hole (Jbramm or port) , or at most a short 

I chink, at the tip. through which the pollen lias in 

J some way to be dischniged. In Vaccininm (Cran- 

m bcrr)-, Bliieborrj", &c.), the pore-bearing tip of the 

yi anther-cell is prolonged considerably, often into a 

]\ slender tube, aa in Fig. 340. In the BarlHTiy (Fig. 

I I 49 J) and 1n most of that family, also in Lauracc». 

„ the whole fcce of each anther-cell separates by a coa- 

linuons line, forming a kind of door, which is attaclied at the top, 

and tnms back, as if on a hinge : in this case, the anthers are said 

to open by uplifled valves. In the Sassafhis and many other 

plants of the Lanrel family, each lol>c of the anther opens by two , 

emallcr valves of the kind, like tra|)-doors. 

401. The attachnieut of the anther to the filament presents 
Uiree prineiiial modes, which ai'c connected by gi'a<latians. 
These are the 

TnnaU (Fig. 49.'i, 49fi), in which the anlhcr directly continitca 
and corresponds to the apex of the filament, the cells usually 
dehiscent strictly marginally, the lobes or cells not looking or 
projecting either inwanl or outwnrtl. 

no. 4». 


FIO. »!. 




no. 4{>». 

mHniuT i1n«i> Il» wlinic lnniOli or tl» nnter Mb uf cacli «ll. 
Stuaenof aPyrolaioKlxwltorUwuilbcropeiil"! hy » tiitBlnal oH- 

Stanum ot a nnrborrr; IIm cell* of the anllior vpenlne owli bjr an up- 


.-ITM, irlrh ailliatii utrnm ■n'lirr. 4M. Slarpun nt <XiioIlcrn 
of jLnnun Cunulsnaa. wlUi adiiau uiUmu uid iirolongad ll|> 


Adnate, in which the connective appears to be a direct con- 
tinuation of the filament, having the anther adherent to the 
anterior or i)osterior face of it, and the lines of dehiscence 
therefore looking inward or outward. Magnolia, Liriodendron 
(Fig. 497), and Asarum (Fig. 499) furnish good examples ; the 
latter conspicuously so, on account of a prominent prolongation 
of the connective or tip of the filament. 

Versatile^ when the anther is attached at some part only of its 
back or front to the tip of the filament, on which in anthesis it 
lightly swings ; as in Plantain, in all Grasses, the Lily, Evening 
Primrose (Fig. 498), &c. 

4()2. The direction to which an anther faces, whether inwaixi 
(toward the centre of the flower), or outward (towaixl the peri- 
anth), has to be considered; except in the case of an innate 
anther with stricth* lateral or marginal dehiscence. An anther is 

Extrorse^ i. e. turned outward, or Posficous^ when it faces to- 
ward the i)erianth, as in Magnolia and Liriodendron (Fig. 497), 
Asanim (Fig. 499), and Iris ; these all being cases of adnate 
and extrorse anthers, the cells attached for their whole length to 
the outside of the summit of the filament or the connective. 

Inlrorse, i. e. turned inwanl, or Anticotis^ when it faces toward 
the axis of the flower; as in N\Tnphieaceae (Fig. 318), in Violet 
and Lobelia (which are adnate and introrse), and in CEnothera. 
In the common Evening Primroses (as in P^ig. 498) the anther 
is fixed near the middle, introrse, and versatile. 

4G3. The direction in which the anther may be said to face, 
outward or inwanl, dei)ends upon two characters, which do not 
always coincide, viz. the insertion or attachment of the cells, 
and the position of their line of dehiscence. In such a strongl}' 
characterized adnate anther as that of Liriodendron (Fig. 497), 
both the attachment and the dehiscence are plainly posticous or 
extrorse : in most sjxjcies of Trillium, the cells are Introrse as to 
attachment, but some are nearly marginal and some are even 
rather extrorse as to dehiscence : in the related Medeola, and in 
Lilium, where the anthers are extrorselv afllxed toward the base 
or middle to a slender tip of the filament, the dehiscence is 
either introrse or nearly marginal. Pamassia is in similar case ; 
the anthers being clearly extrorse as to insertion and more or 
less introrse as to dehiscence. 

4G4. Adnate anthers are iKjrhaps as frequently extrorse as 
introrse. Others, whether basi fixed or medi fixed ^ are more com- 
monlv introrse. Those fixed bv the middle, or at anv other 
part of the back, and lying on the inner side of the filament, 
arc said to be Incumbent, 



4G5. The connective may Iw app<;ni.lageti either by a pW 
gfttion or ollierwisc fl-om tlie tip (as in Pig. 491)), or from tUo 
bac-lc. SB III Violets and in many Ericnccoiis plant*. 

4GC. The nonnal anther is two-eellecl, biloetdar, or (to use 
a )cs9 common term) dilkeeout, and its lobcB or cells |MirH]lt.'l, 
right and left ; but tlie cells at first, and sometimes at inatiiri^, 
are liiloeelUat. that is cacli is di>-iiled into two by a partition 
which stretches IVom the connective to thu sutuix' or line of 

dehiscence. In an 

i) innate anther, and 

nany olhers. thia 

line of dehiscence is 
mi^nal or lateral, 

either strictly or 

nearlj- so, as in 
Fig. 500. When introrsc or extrorsc (as in Fig. 501, 502), 
the sutures may still be considerwl to repn^sent the margina 
turned inward or outward. The pollen is accordinglj- pro 
duccd in four cavities or separate portions of the interior. But 
tlie two lueelli on tlie same side of the midrib or connective 
(riglit and left) are usually confluent into one pollcn-fllletl cavi^- 
or cell at maturity if not earlier, or at least the partition botweon 
them breaks n[i at dehiscence. Sometimes it remains, and, tbe 
groove at the sutures t>eing deep, the anther is strongly four- 
lobed or guadnloetilar at maturity, as in Menis[iermum (Fig. 
504) ; hut morphologically this is still only bilocular (dithecous) 
although ([uadrllwellate. and the anther opens at the sutures 
anil through these pariitiona. 

4G". A stamen being the homologtie of a leaf, the natural 
ffiipixisition is that the anther is homologous with the Wade or 
an a|)ieal iM>rtion of the Made, therefore the two lobes or thecie 
with Ihe right and left halves of it. the int«n'ening connective 
with Ihe midrib, and the line of dehiscence with the lejif-mar- 
gins.' This conception is exemplified l)y the aeeompanjing 

■ Thia n lliv vivnr lunft tu't tukL'ii by Caininj Rtid l(<rpcr, rikI it may »till 
be maintained ii« ttio beil mofphological ™nci'plion. Molil inicrpriBwl wimo 
objcclloaa to il» univeriwllt]' j bnt, at prpnenttil In Sni-'ha'a Teict-Bn»k. they 
are not incnnipatibl^ with tliv conimnn morpholotiy. Saehe takp« the Ala- 
Fnenl witli the cniiDectlve to he the hnmoloEue of the whole leitr, nod Ihe 
•nllipr-cella u appendagei. Olhen. in likening the anlbi>n lo glmid*. adopt 
■ ilmiUr view. 

FtO. IWi, tnimte ant hR. urns aa FIc II 
•>i<i«li>( the CHif laealll. M>l. Samo or an 
SK. SameulliBrrwIlnel-nlniatiman'l 
hjr tbii vulliliini or breaking ur of ifaa luirti 

li •■ Pie. ♦ 



diagram, Fig. 503, which should, however, show the median 

pailitions in the cross-section, or traces of them. Pollen is a 

si^ecial development into peculiar cells of what would 

be parenclnnna in a leaf. Its formation normally 

begins in four places, which may remain separate 

up to maturity', or the two on each side of the axis 

or connective may earl}' be confluent into one cell. 

4G8. Of the manj- de\iations of the topical two- 
celled anther, with its cells parallel and united 
longitudinally by a connective, the simplest and 
commonest is that in which (as in Fig. 505) the 
two cells diverge below and remain united only at 
their apex. Next, the two cells may, in their earl}' 
development, become confluent at the ajxjx, as in the 
Mallow family (Fig. 50G), so as to form a continuous pollin- 
iferous cavity within, opening by a continuous suture round the 
margin : here the anther is unilocular or one-celled by confluence. 
In another wa}', the anthers of some species of Orthocarpus (gen- 
erally resembling Fig. 505, but the lobes or cells quite distinct 
or even separated at apex) lose one of the cells by partial or 
complete non-development and so become one-celled by abor- 
tion. The anther of Gomphrena (Fig. 507) is completely uni- 
locular by abortion or suppression of the companion cell. Thus 
losing one half, it is said to be dimidiate^ or halved. 

461). The two anther-cells, such as those of Fig. 505, some- 
times diverge so much that they form a straight line transverse 
to the filament, as in Mouarda (Fig. 
508) , m which their contiguous ends 
so coalesce as to give the appearance 
of a one-celle<l anther fixed by the 
middle. Or, again, the two cells may 
ha separated by the enlargement of 
the connective between them, as in Cal- 
amintha. Fig. 509. This enlargement 
is extreme in the great genus Salvia, in 
which a very long and narrow connec- 
tive gives the appearance of a filament astride the apex of the 

FTG. 503. Diagram to illastrato the mmrphologj of the utMnen, on the i<1ea that the 
anther answers to leaf>blade: the lower portion being filament and a r*rtof the anther, 
in section, the upper a part of a leaf. 

FTG. SM. Stamen of Menispennnm Canadense, the qnadrilnrellate anther divided. 

FIG. SA9. Stamen of Pentstemon pnbesoens, with antber-celln divergent. 

FTG. 006. Staroenof Mallow (one of the closter of Fig. 486K the two cells and latures 
confluent into one. 

FIG. sm. Anther of GomplinDa or Globe Amaranth, medilixed, of a single cell, 



proper lilumont, ami bearing an aiitlier-coU at oacli end. 

few a{xrc-ie9, the two aether-cells are nearly alike j in more, QuiJ 

loirer one is imperrect, as in Fig. 610*; in more, it is nbortive [ 

or wanting altogetUor. as in Fig. 510*. Then, in the relatt'd j 
Calirornian genus AiidtLiertia, the lower half of this eonneotire i 

is reduced to a short tail, as shown in Fig. 511', or even In 
most i)f Uie species to so minute a vestige that, except for tlieae 
trnnsitinns, the stamen migiit Iw supposed to consist of a simple 
filament, tvitli an interniption like a splice in the middle, and 
surmuiinted liy a one-celled anther, as shown in Fig. 511*. In 
Koscmary. tlie continuity is complete, although a minute reflexed 
tooth sometimes indicates the junction. 

470. l^illeii, the product of the anther, is usually n powder)* 
gubstance. wliicli wlien magnified is seen to consist of separate 
grains, of ilefmite size and shape. uniToi'm in Die same plant, 

hilt often rerj" differenl in <liffercnt species or families. The 
grains arc oummotily single cells, globular or oval in shape, and 
of a yellow i-okir. But in Spidcrwort they are oblong : in t 

I. Aiillinn, vllh nppn lart of Blmmnil. nf htbthI IjiMUm. EM. Of | 
. Of«l.:iliimliitljii. MO Oftwo»i«lM«fSii|vta.«llhlnn(iin<!«taiiito 
I utilor fiirk <i( wlilrh l-nar* in* nnHHT-rFll ; tlw Inwer In a (from Saliti 
iiglhiolhi>r(e1MnanlintiDrrn.-tcnnilltlnn; In Alfmni S. nwrlnnl.liMlv 
. en. t. Samoiir Anitlhenu oriHi'llilorii. Iho Inovr Tiik of thr onnrs- 
■ Mkal •par; I., thm A. Mtehyl-h-: In vlil'-li (1>l> \'-v-r (l.rk It umtlf 
Uiv U|i|i>r In In ■ ntnOgfal line wlUi (lu Bluoeiit wlikb 11 •ecm* ta 

nl, tma MlBulus moBcbaliu i Si3. Skrui Sit, 

POLLEN. 257 

Cichor}' and Thistle tribes, many-sided; in the Musk-plant, 
spirally groove<l ; in the Mallow family and the Squash and 

^ £> ^ 

516 517 518 519 530 

Pumpkin, beset with briskly projections, &c. The pollen of 

Pine, as well as that of the Onagracese, is not so simple, but 

apixjars to consist of three or four blended cells ; 

that of most Ericacece evidently consists of four 

grains or cells united. (Fig. 512-521.) The 

most extraordinary shape is that of Zostera, or 

the Eel-grass of salt-water, in which the grains 

(destitute of the outer coat) consist of long and 

slender threads, which, as the}* lie side by side in the anther, 

resemble a skein of silk. 

471. Pollen-grains are usuall}' formed in fours, by the division 
of the living contents of mother cells first into two, and these 
again into two parts, which become specialized cells. As the 
pollen completes its growth, the walls of the mother cells are 
usually obliterated. But sometimes these cells 
persist, either as shrecls, forming the cobweb-like 
threads mixed with the ix)llen of Evening Primrose, 
or as a kind of tissue combining the pollen into 
coherent masses, of various consistence. Of this 
kind are the elasticall}- coherent pollen-masses (or 
PoLLiNiA, sing. Pollinium) of Orchises (Fig. 463), 
and tlie denser waxy ones of many other orchids and those of 
Asclepias or Milkweed, Fig. 522. 

472. A pollen-grain has two coats. The outer coat is com- 
paratively thick, and often granular or flesh}*. This is later 
formed than the inner, and b}' a kind of secretion from it : to it 
all the markings belong. The inner coat, which is the proper 
cell-wall, is a veiy* thin, delicate, transparent and colorless mem- 
brane, of considerable strength for its thickness. The i>ollen 
of Zostora and of some other aquatic plants is destitute of the 
outer coat. 

473. The cavity enclosed by the coats is filled with a viscid 
substance, which often appears slightl}* turbid under the higher 
powers of onlinar}* microscopes, and, when submitted to a mag- 

FTG. 516-521. Forms of pollen: 516, Lily; 517, Clchory; 618, Pine; 519, Circiea; 
020. Kftlmia; 521. Rvenin^; Primrofle. 

FIG. 522. A pair of iwllinla of Aacleplas, annexed by their candicles to the gland. 




iiifjing power of abotil. three huiiclred (Uametcrs, is Tound to J 
contain a miillitiKle of miniito particles (fuviUa), the larger of I 
wliicli ai'e fhini one four-thousaDtltb to one live-thousandth of an I 
inch in length, and the smaller only one fourth or one sixth of f 
this Bize. When wetted, tlie grains of jwllen promptly imbibo I 
water by endosmosis, and arc distended, changing their shape I 
somewhat, and oMIU'rating the longitudinal folds, one or n 
In number, which many gntins exhibit in the dry state. Soon j 
the more extensible and elastic inner coat inclines to force its \ 
way Ihniugh the weaker parts of the outer, especially at one or j 
more thill points or pores ; sometimes forming projections, 
when the absorjition is slow and the exterior coating tongh. In 
many kinds of [>oUen. the grains, when immersed in water, soon 
distend to imrsting, discharging the contents.' 

474. Pollen-tabes. In others, and in most fh>sb |wtlen. wheo 
placed in oiilinarity aerated water, at least when this is slightly i 
thickened by syrup or the like, and submitted to a congenial teni- 
peratin'e, a projection of the inner coat through the outer appoara 
at some one poinl, and by a kind of germination grows into a 
slender tul*e, which may even attain two or three hundred times . 
tlic diameter of tlie grain ; and the richer jirotoplasmic contents I 
tend to accumulate at the farther and somewhat enlarging ex- 
tremity of this |»ollcn-tubc.' 

47a. In oleistogamous flowers (434), the pollen, while still in \ 
the anther, sends out its tubes, which may grow to a great length, 
in the mere moisture of the flower-bud. the growing tip always J 
directing itself toward the stigma in a wonderful way. Similarly, 
in the o|>en Bower of Milkweeds, the pollen-tubes eomeUmes ] 
start fVom the |M)llen-mass even while yet in the anther, and In 1 
vast numbers, forming a tuft or skein of polIen-tul>cs, which ] 
may attain considerable lengUi and direct itself toward the somo- J 
what distant stigma. Commonly, however, the pollen remaina I 

' In Coniferre, Ihe grains of pollpn Imve s peculiar internal i 
mherB dcrclupmont (•u^gcMlTi of ■ lioinultig}- willi the micrMporM o( « 
of the higher Cryplos»nii<i|, ihc Ponli-nU «l or lieforc malurily ondenpiiac'l 
dlTtoion Inio two or thrw Inlcnwl oclU. only one of wliii* acti in rertUii*' I 
tion. When Ihcy «ci ii(H)n ihe ovuie or are placed in w»ter, and llir innerfl 
co»l •wells liy MlwoqiUun. tlie biirtlioK oiiier ciwl i* voniniunly (hrowo oft ■ 
In Pines nnil Firs (but not in Ijircli am) Hemlock Spruoi'). the grain oCjl 
pollen i« »lngularly enmponnil. ™n»i«tina (as in Fijt. fil8) iif a it'tilml •rcuata f 
body Ithe proper pollen-cclll bearing at cacti eniT an empty roundish cell. I 
Tlieac are »e«icular protrniionii or appeniUpc* of the proper ]H>1len-(Eriiin, ot 1 
no known funclional importance, except lliat they mider auch wiDil-<li»- 1 
perted pollen more buoyant tor tran«iiorTAiirin, 

* Van Ticghcm. in Ann. Sci. Nst. wr. o. %a. 313. &c, IBdO. 


unaltei'ed until it is placed upon the stigma. The more or less 
viscid moisture of this incites a sim- so sh as 

ilar growth, and also doubtless nour- 
ishes it; and the protruding tube at 
once i^enetrates the stigma, and b^' glid- 
ins: between its loose cells buries itself 
in the tissue of the style, descending 
thence to the interior of the ovary and 
at length to the ovules. Fertilization 
is accomplished b}' the action of this 
ix)llen-tube n\x>n the ovule, and upon a 
si>ecial formation within it. Consequent 5» 

upon this an embiyo is formed ; and the ovule now becomes a seed. 

Section VII. The Pistils, ob Gyncecium. 
§ 1. Is Angiosperms. 

476. The succinct description of the pistil in the first section 
of this chapter (302), as also what has been stated of the modi- 
fications of the g3'noecium in Section III., relates to the most 
t^'pical conditions of this part of the fiower. The essential 
characteristics of all ordinary- pistils, whether simple or compound, 
are : 1. a closed ovar^', in which one or more ovules are included ; 
and 2. a stigma, uix)n which pollen for fertilizing the ovules is 
received, and through which the pollen acts upon them. There 
is a more simplifie<l condition, in G3'mnosperms, in which naked 
ovules are exposed to the direct action of the pollen. In con- 
tradistinction to this, the ordinary pistil is said to be Angiosper- 
mous ; that is, with the see<ls enclosed in a sac or covering, this 
in the fiower being the ovar}'.^ And plants with such gjnoecium 
are denominated Angiospeums or Axgiospermous plants. To 
such onh' the present subsection specificall}- relates. 

477. The several terms which apply to the Gynctcium or 
female system of a fiower, and to its components, have been 

^ Although thus originated, the seeds are not in all cases matured in a 
closed pistil. In the Blue Coliosh, Caulophyllum thalictroides, the ovules 
rupture the ovary soon after flowering, and tlie seeds become naked ; and in 
Mignonette they are imperfectly enclosed, the ovary being open at the 
summit from an early perio*! of fructification. 

FIG. ISSZ. A pnllen-grain of D»tara Stnimoniatn. emitting Its tnlie. R24. Pollen- 
grain of a Convolvnla^. with Itii tube 525. Other pnl1en-in*iiint, with their tiibef*. leM 
strongly niagnifletl. 526. A pollen-grmin of the Evening Primmso. renting on a portion 
of the utignia. Into which the tube emitted tmtxi one of the angles penetrates ; the oppo* 
site angle olno emitting a poUen^tabe. All highly magnified. 



eniimc- rated and ilefined already (30*2, note) : the elementary 
term is that nf 

■17^1. Carpel. Lnt. CAKFEtLrM. This is the tei-m coined hv Dttnsl, 
and is in eoinnioii use. The I>ettei-fomied wonl Caih-iihuic 
(English Carpid) has been [jroposed, and best uf all Cabto^J 
FinXLlM, in English Cai-pophyll. For carpels arc, as tile word'l 
caiiioiihjlla denotes, pistil- leaves, or leaves of the gjncecium, 
1. e., seed-bearing or IVuetiferous phylla. Thej- occupy- tlie ocQ- \ 
tral or iiiipermost region of tlic flower. A car|H-l may be a pistil 
of itself, either tlie only one of a blossom or one of several, or 
it may be a eoDstitiienl of a more complex pistil. Ju either case, , 
a carix'l is the bomologue of a leaf. 

47JI. The morphological conception of an imcombined caipel I 
is that of the blade of a leaf incnn'ed lengthwise, so tliati 
the margins meet, and join by a snture, thus furming a closed j 
sac, the oi-an/. A prolongation of the tip of the leaf b thexy/ir.- 
some i»rtion of this, usually the apex, not rnrely a a 
doiiljJe line down the side wliieh answets to the suture of tho I 
leaf-margins, an<l may be regarded as its continuation, is the 'I 
stignm. The carpellary leal' is always tnctin'e<l : the Jower sur- J 
face of the loaf is represented by J 
thu exterior surface of the ovarj', 
the upper by the interior. The 1 
conjuined maj'gins of tlie leaf, or 1 
whatever they l)ear, are intemiil 1 
in the ovary : the stigma may lie I 
rcganled as a jjortion of leof^ j 
margins presented externally , rtc«- 
titnte of epidermis and (brmed 
of loose cellular tissue, which in 
antliesia is moist by some secretion. The onilet are peculiar 
struflnn's normally arising as outgrowths from tlie mai^ins of 
the leaf, or some part of them, eometintes from the whole or 
u special portion of llic upper or inner surface of the leaf. 

480. The car|K'llar)- leaf l>eing involute, the suture, on vhkA j 
the ovules are normally borne, always looks townnl the axis OF 1 
centix< of tho Hower. It is the only proi>er suture (or sesin) e f 
carix'l can have. Prom its |)osition it takes the name of / 
or Ventral SHlurv. Ami the opijosit*- line or riilge, answerJDg I 
to the midi-ib of the leaf, being sometimes prominent ami of tlie ' 

Flo. K!7. A 1«r IncnrtlnR, to llluxnl- Iks ninrpli.ilni^ nl»ttnivi« r>l>tll nr CMpri- j 

IBM, A mti»t(nf Impynim bllwtinlninl. rnl - . . ...•.—- 

• itnnlili' Uh>I unit tlw imlBr" bwirlni the ••< 
rnti-i It AUnb M>rUM.| wlikli liu nponnl 


api^arance of a suture, has been somewhat incouginiously named 
the Outer or Dorsal Suture, 

48 1 . The number of carpels in a g3'noecium is simpl}' expressed 
b}- adjective terms consisting of Greek numerals prefixed to this 
word : e. g., Monocarpellary^ of a solitaiy carpel ; DicarpeUartj^ of 
two carpels ; Tricarpellary^ of three ; Tetracarpellary^ of four ; 
Pentacarpellary^ of five, and so on up to Polycarpellary^ of many 
or at least of several and an indefinite number. Less general 
and on I}' i>artialh' s3'non3'mous terms are such as Mouogymus 
(of one pistil), Diyynous (of two), Polygytwus (of man}'), &c. 
These are adjective foims of the names of the oixiers, from 
Monoyynia to Polyyynia^ in the Linnaean artificial classification, 
which either supposes the caqxjls to be separate or parti}* so, or 
confounds simple and compound pistils. 

482. When the g}noecium is of a sohtary carpel, the position 
of this as regards the axis of inflorescence is not uniform ; but 
commonly its back or dorsal suture is before the subtending 
bract, or in other words the ventral or ovule-bearing sutui-e 
faces the axis of inflorescence. When there are two carpels, 
they face each other, bringing their ventral sutures into opposi- 
tion, and as to axis of inflorescence either median or transverse 
(291), but usuall}' median, that is antero-posterior or in the 
line of bract and axis. Cruciferaj, Capparidacece, and Fumari- 
aceae are somewhat remarkable for having their two carpels 
right and led, that is, collateral or, in other wonls, transverse. 
When three, four, or a greater number, the}' divide the circle 
equally, or when numerous they take a spiral instead of verticil- 
late onler, and occupy several or many ranks, as in Ranunculus, 
Magnolia, Potentilla, &c. 

483. The Gynoecium may be either of separate carpels 
(A/>ocarpous) , or of carpels coalescent into one body (Syncar' 

pons), or of all grades between the two. Apocaqx)us pistils are 
sinij)fe ; a 8yncari)OUS pistil is compound, 

484. In both, the essential parts are the ovary and the stigma. 
The style may Ix; conspicuous and widely separate these two, 
as in Fig. 530-538 ; or haixily any, as in Fig. 532-535 ; or none 
at all, as in Fig. 530, 531, 533. 

485. Placenta. This name' is applied to any surface in the 
intenor of the ovar\' on which ovules are borne. It has been 
stated (570) that these are usually lx)rne ui)on the mai*gins of 

^ Taken from a remote analogy with the placenta of the lii|;Iu>r animals. 
The name appears to have been introduced into Iwtany by Adanson It has 
been termed Trophosftermum or Spermophorum by some of the early modern 


<^r|ifllan' leaf, or apon some portion of whst t 

AVIicn the dvules are niiaieroii.s, and t 
\ tinics when tlioy are few, the coiuhiiiorl leaf-trct^c* 1 
triilarge to form a kind of receptacle for their attactH I 
mt-nt or supjKul : this is the Placenta. In Fig. 530, I 
the plnceuta ie well developed, and also ia Bucb | 
ttynoarifOUB ovaries as arc illustrated in Fig. 536. 
'', 544, and 545. In vcrj' many others (aucli as 
Fig. 52K, 531, 533), there is uo particular eolai^gv^ | 
ment of the leaf-mai^ns visible, and no jiaiticular 1 
ground for the use of this epecial tcrrn. 8UU it I 
is commonly us«i, ns occasion serves, even for tbe ( 
mere line or spot on which ovules ore borne, i 
wi'll as for a more promiDent development to wbicb 
ia> llic name wns originally apjilicd. 

480. Hlmple or Aporarpons PMIIfl mny be solitarj. several, or 
nnraemn*. When indctliiili-Iy numcroiii^. they arc seldom in one 
ciiflc. but arc eopitatc or spicate u|>on a proportion iitcly enlarged 
or |)rolonged receptacle, as in Anemone, Rauunculus. and most 
ntrikingly in Myofliims ; when rtKluccd to a single one, as in ' 
Actwa, I'odopliyllum,' liorbcrrj'. and Plum or Cherry, the car- | 
[lel inoNtly api^ears oa if it were an actual termination of the i 
floral axis. But even then the pistU i 
is hnrdlj- ever quite symmetricod in 
sliaiic : the ovary is somewhat gib- i 
bous or unequal -sided (as in Fig, 
312,315,316,528,531-533), and Uie J 
stigma more or less oblique or avca | 
wholly lateral. The continuation of i 
the latter down the whole length of 
the ventral aide of the style (as in Fig. 5i**. and also Fig. 549) 
is not uncommon. In Scliizandra (Fig. 531) it is continued i 
diiwnwnrd on thu ventral edge of Ihc ovary as far as to ita i 

■ Abnormal ipedmcni of PodophjHuro pchntura are occRtionslly found I 
ImviiiK a gj'niH.-lum of from iwo to lix ■rponilc tarpeU. 

* I'lruriigyne, ■ Orntlanactou* tceniu *o name*) on thii account, ha 
MjXir nor apical >tl|tnia wliati>Ter, but ha> a lung ttifnna pxlciidinnc down Ihe 1 
uulklilt' ul Mu:li oitulili'mus auturv of ill lUcarpellaty ovary lor mogt of its f 

FIO. SSO. alnCloilniplaiilHllaf Poiliiplijlluin.cut KKiHtii*bnwlbe|itac*nla,a 
PIU. mi. VarlloLl HcUon uT ■ i^ntll of Si'lilumln nicrlnu; ■ ililii vtrw diowlnc ' 
tlK>UcinitilnuTT«nt<b><mloIl>emlilrl1e<>tlliraTir;, Wi DXilnrtlylnwtia: n " " ' 
tl'w. MX rtaUlof Amawrabn,DHtaerflBi,Bi aa Wibow Vha Inlarliir of Uia o\ 


487. As the placenta of a simple pistil belongs to the two 
united margins of the caqxjllarj' leaf, there is naturall}' a double 
row of ovules, one to each margin. If the leaf- 
margins which are turned inward in the ovarj' be- 
low to bear the ovules are turned outward above to 
receive the pollen (see Fig. 531), then the tjpical 
stigma should also be double or bilamellar. So it 
is seen to be in such cari)els as those of Fig. 528, 
531-533, and indeed in ver}' many stigmas of this 
class. Such division, or even a greater bifurcation 
of a monocar^xjllar^' stigma into two lobes or half- 
stigmas, is not anomalous. 

4^. The ovary of a sunple pistil should be 
unilocular^ that is, should have a single cavity 
or cell (loculus), although, as will soon be seen, 
the converse does not hold true. Yet this cell in 
certain instances becomes hilocellcUe^ being divided 
by a growth or intrusion from the back into two 
locelli. This occurs more or less in the larger 
number of species of the Leguminous genus 
Astragalus, and the mode is shown in Fig. 534. 

489. Compoand or Sjncarpoiis Pistil.^ This consists of two, 
three, or a greater number of carpels coalescent into one bod^*. 
A tnie compound pistil represents a whorl (in the simplest case 
a pair) of carpels united into one body, at least as to the ovar}'. 

490. The coalescence of a capitate or spicate mass of carpels 
or simple pistils of the same flower, imbricately hcai^ed on the 
torus, as in Magnolia (Fig. 648) and Liriodendron, cannot 
projierly be said to form a compound pistil. This heap of 
pistils may be called a Sorema. 

491. Moiphologically, a compound pistil, as to the ovaiy, may 
be a j)air or a circle of closed carpels or simple pistils brought 
into contact, and the contiguous parts united : this is illustrate<l 
in Fig. 535-538. Or it may be formed of a whorl of o|)en car- 
jxillar}' leaves, joined each to each by the contiguous margins, 

1 Tlie terms ajiorarpoug and syncarpous for pistils, tlie first of separate, tlie 
second of combined carpels, were introduced by Lindley. They have little 
advantage over the terms simple and compound. Moreover, the word 
tynctirp or xffnrarpium had been appropriate<l to a sort of fruit of the class 
now calk^l multiple, formed by the coalescence of several flowers, and also 
to that of a heap, head, or spike of carpels more or less cohering at matu- 
rity, as in a blackberry, or confluent in the flower, as in Magnolia. 

FIG. .'S.'U. Ovary or fnrminK legnme of Astragalas CanadenMn. transvenely divided, 
to Rhow the falne i»artiUou wlikb, intmded from the back, divides the simple cell into 
two half-cells or tocelii. 



in the manner of Fig, 542-i>45. Between tliese two Uiei^ 
evcrj- grailutioii. The fiist fomiB a coiiiiMund ovar\-. 

1:12. Wllh Iwo or more Cells iind AxIIc Placentie. For it is 
cvMeiil that, if the contiguous iiurts of a wlioil of two or more 
closed cariwla eoliere, the resulting compound ovnry alionld liave 
OH many ccUu as Uiere arc cai-pels in iu compo§ition. nnd tluU 
Uiu i)l«ct-iita- (one in tlie inner angle of each carpel) will all be 

brought together in the axis of the compound pistil. And the 
partitions, termed Dissepiments, which divide the corajwund 
ovnry into cellq, manifestly consist of the imitecl contiguous por- 
tions of the walls of tiio caqtels. Tliese necessarily' are composed 
of two layers, one belonging to each carpel ; and in fHiit Ihey 
often split into the two lajera. Tnie dissepiments and the tme 
cells must accordingly 1>c equal in numhcr to the cai-jiela of 
which the compound pistil is comi>osed. That is. the ovarj-, or 
the resulting fruit, is bilocular or 2-oelled, trlloeutar or S-ccIled, 
if»adriloeal/ir or 4-celled, and so on. according to the uumber of 
dissepiments or cell 8. 

493. There may also be falte diueprmmit, mostly of Uio same 
ciioraotcr as that which In Fig. 534 divides the cell of a single 
carpel. Such ore found in Fhix (Fig. 539-511), in Amdanchier 
or Senice-ltcrn% in Unckleberrj- (Gaylussacia), and in moat of 

riG, ma. PiMll of k Saximci! trunpawl nf iwo ciir|«1> 
bakxr. Iml illMlnirl abuvu^ nil unw both Hbnre •n.1 belov. 

ria. U6. Plilll of comnuRi St. Johnnurt, of Uim anltvl <»arl«: 

ilmpls pIMIl* nnlt^ 

ria. m The r 

le of unothsr ipKlc* of Bt. 
Inln nna, wliloh. hnwew, mi 
'Tnulnmlla "t Splrtcrworl 


tiic American species of Vaccinium. In all these, the false par- 
tition [9 a growth from the middle of the back of eaoU carpel, 
which divides its cell more or less completely into two. 

494. On the other hand, even the troe dissepiraents which 
belong to such a. compound ovarj- may be abortive or evanescent, 
the placenta remaining in the axis combined into a column. 
(49£l.) The second modification of the compound pistil (491) 
normally has an ovarj', 

49a. With one Cell and Parietal Placentn. That is, the 
placentic are borne (as the tenn denotes) on the waQ or |>arietes 
of the ovnry, as in the Poppy, Violet, Sundew, 
Cistus or Helianthemum (Fig. 543), Cleome, Gen- 
tian, and in all or most of the orders fVom which 
these osampks are cited. The diagram Fig. 642 
, illustrates the morphological conception of a com- 
' i>ound pistil of tliis kind. Not that it is ever 8ii|>- 
lH)8ed to be formed by the actual combination of once 

separate leaves, any more than a gamopbyllous calyx or corolla 
is actually so protliiced. The conception in all such cases is that 

FIG. 330. TnDircTM dlogncuDiallc MClton oft flower of the <»inmon Flax, sfanw- 
InK ILc rivarj irlth hlw imrtttlnm eiteniilng nne from the back of eactl i-ell. OtO. Sec- 
tion of aniiUaie fruit >ml imls of the uime. tlie blM! pari It Ion* now comgilple. .llvM- 
lliK the flee islle Into ten. oub nno-Kedeil, Ml. Same ota wUJ FIbje iLliiuui |«TCDne), 
111 wlikh the taiKO pantMiinii n-m:iiii tiicnnipl»l«. 

FKI. SU. Pbui nf a nnMxtleil nvary villi Ilirce parietal flacentr. rut nrrnw he- 
iim; Die upper part diowbig the (up gf the three leaven it In thmrcUcHllf comjiuBed of, 

Fid. M3. Ovuj of HcUauthemDin Canadenu, cat aemw. ihuiriiig the mule* on 
thrcv (larivlal placcntn. 

Fir,. IH. T™i«*rM»«;tlnnoftheo»ar,»ofH}'perl<-ninmiv6..1en<; lbs three larfte 
Iilai-eniai nieelliigln tbecenlie, but nati»beriiiK. US. Sliullai KCtlou of a ripe capaida 
oftheaame; Uieplacenlie now avldentlT pwlctol. 



of a coDgenilnl development of organs in union nbich, in the I 
develo|niient of a vegetative sLoot, wotilil 1)e leaves. This c 
is I'eiJRiBeiii^d bv llie c-oiiil>iuqtioii of o[ien carpellary leaies. »a 
lUe |ireoe<iiiig one is by that of closwl ones. As the etigoa 
of tbe leaves iuu8t neeils be tunied in, to bear tlie ovules, 
a comixiund ovaiy with parietal plneentntion may be lilvcm-'d 
to tlie unopened calyx of a Clemulis. as Bliumi in Pig. 2oG, 
257. Eier)- gradation is found between asile and lutrielal 
plftcentaliou. Sometimes the plaeeutK are strictly oti tbe pori- 
ctes or wall (Fig- 513, 547) ; sometimes Itonie iiiwnixls on 
incomplete Uiasepimeiits (Fig. 648) ; and sometimes tliey arc 
brought firmly together in tbe axis, as in Fig. 544, though seiui- 
rable, and indeed separated in the fi-tiiting stage. 

496. A compomid ovaiy with parietal placentte is necessarily 
one-celled {uniloeu/ar) ; except it be divideil bv an anomalous 
partition, such as is found in Cruoii'erie (Fig. S'-io) and in many 

4a7, Normal placentte are necessaiily double : when jiarietal. 
the two halves lielung to diflerent leaves ; when asile, to tlic same 
leaf. These two halves may diverge or l>e widely separated, 

sometimes even at their oiigin. as in Aphyllon anil some other 
Oro ban clia cere, in which a lilcaniellary ovary has four almost 
equidistant placenta; ; or in hucIi cases the placcutre may be 
regaitled as tntra-mai^nal instead of mai^nal. 

4!t8. The plncentje of a two-sevcral-eelled ovar^-. such as in 
Fig. 53G, 5.17, ic, may be described In the plural number, 
being one in each carpel ; or when consolidated uilo a central 
column, and well covered with ovules, they may be said to form 
one (com|>oiiiid) [ilaeenln. Then wlien tlie dissefiiments early 
disappear, or arc abortive fVom tlie first, tbe result is a compoond 
ovary of this class. 

4'Jll. With one Cell and FrM Central Placenta. In Caryo- 
phj'llnceic (Fig. 54!), 5,iO) and Portulacaeete. this evidently 
results from the obliteration of the dissepiments (as many as 
there are styles or stigmas) . vestiges of which may be sometimes 

Pin. MC IHiiRniiii (er>niiiil>rliin > In niarlnto rwv nninil iilnamiatlnii rmatan^ 
l<*itlii>nt<ninr-llii>rplmniu. MT. Ssine>rr«trlFi |Hir1Mitl|ilw«nuniin. Mti. Saain i 
Ills |>UcvuI]e canlfll luWBnl on Imlistlecl dlacjilmoDU. 


detected, while certain plants of the same families, of otherwise 
identical structure, retain the dissepiments even in the fruit. 

500. But a similar condition ma}- equall}' ailse 

from a modification of parietal placentation, namely, ^ 

with the margins of the leaves ovuhferous only at 
bottom, and the placentae there conspicuously devel- 
oi)ed and completely* united. The basal placenta- 
tion of Diona?a is unavoidably- so explained, its 
nearest relative, Drosera (Fig. 553), having parietal 
placenta?. And this leads to a probable explanation 
of the case in Primulaceoe, where a large free central 
placenta fills the centre of the cell, and no trace of 
dissepiments can be detected.* 

501. The idea maintained in former editions is 
still adhered to; namel}', that pkcenta? belong to 
carpels and not to the cauline axis, in other words, "** 
that ovules are productions of and borne upon leaves, usually 
upon their margins, not veiy rarel}' u|X)n other ix)rtion8 of their 
upper surface, rarcl}' over the whole of it.^ 

502. Ovules cover the whole internal face of the carpels in 
Butomus and its relatives, also of the Water-Lilies (both N^m- 
pho^a and Xuphar, Fig. 551) excepting the inner angle, to which 
they are usually restricted in other plants. And in the allied 
Brasenia and Cabomba, where the o\'ules are reduced to two or 
three, one or more of them is on the midrib, but none on the 

1 The placenta in this and like cases is rather to be regarded as an out- 
gn)wtli from the base of the carpellary leaves, conibiniHl over the floral 
axi.s. Upon this inter])r(^tation, a central portion of the column may be 
(and sometimes must be) of axile nature, yet the ovules be borne upon 
foliar parts. See Van Tieghem, in Ann. Sci. Nat. ser. 5, xii. 320 (1800) ; 
Celakowsky, Vergleichende Darstellung der Placenten, &c. (1870) ; Warming, 
in Ann. Sci. Nat. ser. 0, v. 102. 

2 This view was first maintained as a general theory, and on critical 
grounds, by Brown, in Plantae Javanica Rariores, 107-112. Schleiden, End- 
licher, and others took the opposite view, i. e., that ovules are productions of 
the axis, even in parietal placentation, — an exceedingly far-fetched suppo- 
sition. In later days, the commoner view has regarded ovules as of both 
origins, as productions of the carpels in parietal, of the axis in at least 
some free central or basilar placentation. But at present the theory of 
foliar origin without exception, revindicated by Van Tieghem, and csjk*- 
cially by Celakowsky and Warming, again prevails. For the bibliography 
and an abstract of the various views, see Eichler, Bliithendiagramme, espe- 
cially the note in the preface to the second part (where he gives his entire 
adhesion to this conclusion) ; also Warming's memoir, De I'Ovule, in Ann. 
Sci. Nat. ser. 0, v. 1877-78. 

FIG. 540. Vertical section thmnfd) tlie coraponnil tricarpellary ovary of Spergularia 
mbra, showing the free central placenta. 660. Transverse section of the same. 


a of tbe cupdJuy kaC In amay apeow of G^ntiaa, wi 
also inObobmsad Buvnia. ortlwHine 
&milj, ifae wbde inttrD*] fiuv of a iUckt- 
peDarv ovarr is tbkUr (Mrulifiitws. 

503. I^riia(is the porictHl I'Ucrntie in 

Psnu^sia (F^. ^i) a» horue on the 

midrilH of the carpels, tc€ Ihcy air iliiwrtl^- 

uDiler the Etigmaa. instead o( attvraate 

iriUi tbem. as Chej oormaOv sboukl be. The eanw tbing ocean 

iu I^>i>pii-s and manr olber P«pavw»c«B. also in sock- Cnicifene ; 

and in some uf tbe i-ascs cscfa stigma 

»^ b more or Irsa two-lobed. This sug- 

i \ gesU the esi^nation,' here profaaUj- ' 

/ \ the true one, which enppoees that the 

/ 1 [itaceutse are Inme un tbe leaf-li 

^^^^ ^M in tbe uonual way. I>ut thai each 

/■j^^S^ stigma is two-parted (as if the car|iel- 

t ^Hg j Inn- leaf were deeply notched at the 

ai>ex, and ao its two slignuttic leaf- ^*!5^ 
margins seiiaratc, as Droscra itlus- .^f^^ 
trotes, fig. 553), and that tbe two \ /^ 
half-stigmaa of adjaivnt carpels ha?e mi 

coalesced into one ImxIv. which wonid 
of course stand over the parietal placenta beneath. Eat-h stigma 
in such a case, as well as each ]>ariclal jilacenta, would consist 
of the united margins of two a^jaoent carpels. 

S 2. Is GvM.vusPEiiMa. 

504, G r»>~09PEKUous (that is, naked-seeded) plants are so 
named liecanse tbe ovules, or Ixniies which are to becuine seeds, 
am fertilized liy direct a|ipliuation of the pollen, which reaches 
and acta npon the nucleus of the ovule itself, not through the 
mHlialion of stigma and style. In Ihe structure of their flowers, 
these plants arc of a low or simplified type, in some respects not 
ob\-iously humolc^ons with tbe Angiosi^erros which now consti- 
tute the immense majority of plitenognmous pbnts. But, up to 
B comiiarativoly late geot<^eal |>i.'i-i(Hl. Gymnosperms np|>ear to 
have been the only flower-bearing plants. Tboy are rei>re8cnt«d 

> Glren h; Drown. In the Plant* Javinlno ItRriorpi, bIhivp reffrnnl to. 
Flo. 0.11. TnuuTrnn hkUihi ufftn ofvyiif MynplumedurftU, Uiiicar|H>UarBllfkr- 


in the extant vegetable kingdom b}' three (or four) groups or 
orders, two of them small, and one eomparativel}' ample and of 
wide distribution ; and all are so strikingly difTei^ent from each 
other tliat the}' cannot be illustrated b}- a common description. 
The largest order, Conifene, is familiar, and contains a good 
share of the most important forest trees of temperate climates. 
The smallest, Gnetaceie, chiefly tropical or of warm regions, 
lies between G^Tunosperms and common Dicot3'ledons. The 
third, C^'cadaceae, is most remote from them, and as much so 
from Monocotyledons, except that it imitates Palms, as it 
also does the Tree-Fems, in habit, both as to stem and foli- 
age. The particular moqihology of G3*mnosperm8 would re- 
quire for its illustration copious details and the history' of various 
conflicting hypotlieses. It must be relegated to the special 
morphology of the natural orders, premising, however, a brief 
sketch of the general floral structure.^ 

505. In GnetaceaB, Gymnosperms and Angiosperms almost 
come together. The flowers have a penanth (diphyllous or 
tctraphy lions) ; the stamens have a distinct filament and anther ; 
and the gynoecium is a sac (presumably of two caipopliylls) 
open at the top and filled at bottom b}- a single ovule of the 
simplest kind, i. e, consisting of a nucleus destitute of coats. 
This pistillar}- body is attenuated and prolonged above the ovule 
into a style-shaped tube, with open and commonl}' two-clefl 
orifice. In the almost hermaphrodite sterile flower of Welwitscliia, 
this takes tlic form of a much dilated stigma, which is even beset 
with seeming stigmatic papillae. If onl^' the pollen were here 
to grow forth into pollen-tubes (with or without a closing of the 
tube) , angiospermy would be attained. But, in fact, the pollen- 
grains bodily reach the ovule itself through the tube, fertilizing 
it directly.^ This interesting group of plants consists of the 

1 References to the literature of gymnospermy and to the steps of the 
prolonged controversy over it, also the points of morphology still in part 
unsettlefl, need not here be given. The history and the idea of gymnospermy 
began with Kobert Brown's paper on Kingia, " with Observations .... on 
the Female Flower of CycadesB and Coniferae," read before the Linnean 
Society in the year 1825, and published in King's Voyage in 1827 ; and the 
bibliography down to a recent date is given by Eichler in Flora Brasiliensis, 
Gymnospermia, iv. 4*^, and in Bliithendiagramme, i. 55-00; also ii. preface x. 
See also Alph. DeCandolle, Prodr. xvi.^ 346, 624. In this volume, the late 
Prof. Parlatore adhered to the ancient ideas in his monograph of the Coniferae. 

2 The view here implicitly adopted is that of Beccari, founded on the study 
of Gnetum, and published in Nuovo Giomale Botanico Italiano, ix. 1877. 
It was before nearly or quite reached in successive steps, by J. D. Hooker, 
in his classical memoir on Welwitschia, in Trans. Linn. Soc. xxiv.; Stras- 
burger, Die Coniferen und die Gnetaceen, 1872; and W. R. McNab, in 
Trans. Linn. Soc. xxrUi. 1872. 



gemis Gnetum, shrubs or trees, willi nearly ILe uapeet of 
ADgiosperma, having broful and pin note ly- veined leaves ; Wel- 
witflL-liia of tropical W. Africa, remarkable for its peraistent 
cotyledons which form the only foliage of a woody and long- 
enduring plant, and for its stem or ti-unk wliich broadens wiUl- 
out lengthening, except in it^ Ho wer-a talks ; also Ephedra, of 
much branched shrubs, inoiuly of wann-t<?mperale regions, lealless 
or nearly so, one species of which inhabits Europe and two the 
southern borders of the United States. 

606. The flowers in all Gymni>si>erms are diclinous, either 
dicGcioiis or moncccioiis ; except that those of the strange G neta- 
ceoiis genus Weliritsthia are stnictumlly polygamous, the male 
flowers hating a well formed but sterile gynoicium. 

'>UT In Conlferv, the largest and most important type, are 
embraced ill the famihar (iymnosperms of temperate regions, 
I*ines, Firs, Cedars, Cj-presses, which 
bcir their flowers in catkin- like ehisters / 
and their fruit in cones, and also the \ 
icws and allied trees which do not^C 
produce cones. Perianth being want- "* 
( mg and the sexes wholl}- separate, the floral type 
1 IS so degraded that it becomes doubtful whether 
I each (luster of anthers, or of ovniifcrous seaica 
3\ ulcs, constitutes a blossom or an inflorcs- 
«. Certain Iwtanists look upon a whole 
catkin and otliers upon a male catkin only, of 
I I*ine or Fir as forming one flower. Jt is here 
Lssumed that tineh stamen 
of the one and each ovn- 
hferous scale of the other 1 
answers to a flower of the ' 
simplest sort.' The anthers 
are extrorse, the cells or 
pollen-sacs belonging to the outer or lower side of a scale or a 

1 It will be Brcn tint, fur the fpmali' Auvrcn. tlif* faltows ot rnun* from 
generally »C(*pted *iew ; Bfici. where llils ii coneetkil, nnnlo^ niRy cxlnid it 
lo the msle cnTkini >U»: ycl in lueh cant, where all the phjlta nf nn Indcflnile 
simple >xia are atsnieiii. apirnlly amiii^l on it, the liiffeiencx; belwcca 
Inltnrearence *nil nisli- ttuwvr cnnipklelj Tiiiisheo. 




F«iM«1o flower of 






tni III m IWnsle 

r llHt 





■HlUIUo o 










connective : Bometimea tlieso sacs or cells are two, and the oi^d 
eviiiently homol(^u8 with an onlinarj- stamen : often they are 
more numerous (from ,,^;t=^ f 1~^'"t*^ /^i^ 
tbwc to twenty) and /^^^?^%l 1/ , \jM y i 
variously- diaposetl. // t, i ft ^W*' 'kJt \ \ 

508. The Yew Fam- 1 
ily (Tasinere) is next to '' 
(inetncere in stnictiire. 
It is gcnerallyrankeil as 
a subonlerof ConifcTiE, _ 

but it inny claim to l>o a distinct order. Ttie gj-na>cium is a 
naked onde, terminnting a stem,' and surrouuded by several 
bracts. After fertilization, an outgrowth of the 
receptacle (or a kind of disk, 4114) makes its 
api>curancc as a ring ginUug its base : this 
grown in height and tluckness, and becomes a 
soft-Beshy cup, imitating a hollow berry, in 
the bottom of which the stony-coated seed 
nestles. (Fig. 434-457.) Very simikir is the 
gyiiojciiim of Torrcya, except that the cii|>- 
shaped disk develops almost simultaneously 
with the ovules, and as it grows becomes aduatc 
to the large seed in the form of a 
fleshy coating. In the Gingko, two 
or more similar omles are nakctlty 
develoi>ed <in a nake<l )H>dunclc, im- 
accompaniwl even by a bract (Fig. 
5^K), and one or more of these ripens 
into the l>em-liko see^l, Fig. 550. 
In I'o<locari>iis there are some sub- 
tending bnicts, and the naked ovule 

• H lilies not therefore follow that the otuIc i» a p«n of the axis, or ii 
tcrniinal in the sense of being its direct contintiaiion. In this rcRan] it may 
be only what llie pi»lil of a Cherry is, whlth lo all appearance is equally a 
terminal pnMliittion, hut is really the reprcM-nt alive of the last leaf of the 
axi*. If eo, thai leaf jg here suppressed to the ulmosl, and replaced by 
what U ordinarily ils out|tn>wih, the ovular nucleus and iu coaL The 
BttTji'lure of Toducarpiu favors Ihie iDterpreialion. 

Fin. .Wg. FdnaloHnwennrOlnaknbllnbaorSalMnirlniunsn'irnlta. SM.. Tort Inn 
of IlK nme enlarRnl. Aner Stnulinnrer. W6- A •trii|nreniii (Hit of the ■■nie. In 
TerlloR] Hrtion. enhil.llInK Ilie mntiin rllrii wlihh forniK tlic Heih. tlia rruiit^cvonKKwl- 

secii. Aflrr I>M-i>liine. 

FIG. .-wn. Fom>i1« flnwerot Pmnrarpnudin orala InvarUdnnacolunin orelavMal 
snii]>irl). nablni'lixl tij bracln. Afler Elcliler. 

Flli, {Mil. MagnlDsil vrttlcal lecllao of ■ ilnilUr llnwsr of PoilocariHu. Attar 



is inFert«d on a more or l«ss It^ngthenetl and fltout support, whidi 
is conceived lo represent the carpel. (Fig, 5G0, 661.) 

509. In the true Conifcnt, to whieli Pines, Cypresses, and all 
sucli cone-tVuiting trees belong, the ovules are borne on or in 
tbe axils of scales which are imbriealcd on a simple axis, in a 
spiuite or eapttate manner ; and the male flowers, each a single 
stamen, are also similarly- spicate or capitate. Botli are com- 
monly ternu^l ament« or ctttkiiis ; and the female 
ones pi-operly so, according to tlio preaent view ; 
but the only scales or the male catkins are [utrts 
of tiie anther, Iwing a dilated tip of tlie connective 
in Hnos, and a scale bearing anther-colls or [lollen- 
*" saca on its back in Cypress, 

rdO, III the Pine tribe tbe flowering female eatkin consists of 
bracts, spiially imbricated on the cauHiie axis: in the usil of 
each bract or sterile scale is developed a scale 
which Ittsara two ovulca, and is therefore regarded 
as of carpellarj' nature. These ovules ar« pro- 
duced on the lower part of the upper face of this 
can)eUary scale, and are wholly adhcn^nt to if 
quite to the orifice, which is directed downwanl. 
(Fig. .502, .563.) The ovnliferows scale In 
l)econiiiig fruetiferons usually much and soon oat- 
grows the bract, which is concealed in the Pine- 
cone (or sometimes obliterate<I) ; but it remains 
conspicuons in sundry Fir-con^s. Aft^r fertil- 
ization, the scales, successively covering each 
otlicr in close imbrication, protect the growing 
™ seeds as elTectnally as would a dosed ovary. 

Sooner or later after ripening the scales divei^e. and the seeds 
peel off the face of the scale with a wing attached, and fall or 
are dispersed by the wind.' 

I Among IhoK who admit *• welt u tlioac who reject gyninospcrmjr, 
there ha« been much conlrovpniy oter the morpholany ot the parlt. With 
the tomier. (h« rliacunfon turns on the chnrACIi^r ot the orutifcnitu ccslv. 
A» to thia, Ihe hypolhfiii oritrinally propoiil hj Mohl. and Hilopinl by 
Hraun. ii now mM to be utiafactnrll)' tIrman«ir«IPd liy Ricniel, in Not. Act 
Nat- Ciir. xxxriji, 1ST0. Sec noir bf EnKPlmaiin in Amer. Jniir. Rci. Dtc 
ISTfl, and aluo thr preface to the teennd pari of Elchler'i Biatheniliaenumne, 

Fin anz. VUwiirtbiup*w(kcenraniiri>rn*r|r"«lo<ifiit:aivb,iluiir1ii(ilia[iaIr 
of aiinal* (nnlt« 

no. on. Slntlar tIsw of ■ nuj«i][uy xtaIh oT a Unh, an^ of a bnct t«1iln.l II. 
SM. Otouiul rlan at tlit lainr In iliacniin. re>er»>l : tbe ui'iwr Dinini .luni.tlns Iha ailt 

dn vm*, th^UnrmriiiK bract. Iha mliLllvon 
III lu IMa. Altai Elcblcl. 


511. In the Araucarin tribe the ovuliferous or carpel-scale is 
throughout smaller than the bract, and is completely adnate to 
it, or nith only the tip free ; that of Araucaria (Fig. AGo) bears 
only one ovule, high on the carpel, the orifice downward as in 
the Fine tribe. In Taxodium, Sequoia, and the like, the cone- 
scale is equally inferred to be composed of bract and carj)el-scale 
united ; and indications of this compositiou are to be obsen-ed. 
The ovules (from two to several) are at the base of tlic scale, 
erect and fVcc. The cone-scales are alternate and siiiral on tlie 
axis, but indistinctly so in Taxodium, the BakI Cypress or 
so-calletl Cypress of the Southern United States. 

512. In tlie true Cypress tribe (Cupressineie) the cone-scales, 
which are never numerons, arc opposite or verticillate, t. «. like 
the foliage- leaves, in whorls of twos, threes, or 
sometimes fours ; and the ovules are from two to 

1878, when it ia futty adnptiHl. It wai Bugfrestcd by cprlain ratlicr common 
tnonstro8tt<o», «nJ by the iwo combined leaves of Sciadopilys. 

Awonlinf to tliin view, the orutifcnms scale in tlic Pine tribe is com- 
posinl of tvo leaves of an arreited and Iransformeil bnineli from Ibc Hxil of 
tlie bract, wliii'li arc in tlic normal manner Iransveree lu tbo lubtcniling 
braci, (irc bore carpcllary, each bearing an ovule on Ibc dort-il face ; the two 
are coalesccnt into one by Ihc union of their posterior eilties, ami ihe «i«le 
tlius formed U llnis developod with dor»al face pre»entc<l ro tlie axii of Hie 
cone, the vcnlnil to the bract. It U therefore a compound open carpel, 
compnrcil of two enrpophjlU. This character of being fructifenius on the 
Iwck or lower side of the leaf occurs in no other phKnogamoiis plants, but {« 
the rule in Fem«, from something like which Conifera; may be supposeil to 
bare been derived ; the ovules of the oik in this regard corresponding to llio 
sporanfrin of the other, 


FIG. .1 


;tl™ (In iliagramlof a bract, adnate carpel-scala. sn.i adnalfl 
^ativ. Attn ElcLhsr. 

ui Arbm^VllB. comldemhly Unrer tlian In na- 
One nf Iho Kal«s remiivi'il hiiiI mnre enlarged, 

(0 ahoiT tbe ulustor of ovulo* luidet It. 


several at or on the haae of each eone-seale, always with ori 
upward. Arbor-ViUe (Fig. SGG, 307) has a single pair of ovut«e 
to the scale ; Jiinii>ers, sometimes only one ; trne Cypresses {as 
in Fig. 5CS), often a dozen or more. At flowering time, the c-one- 
scales mostly apijear as if simple ; but in most genera they soon 
thicken greatly within i and they are usually understood to be 
comiJOscd of hraet and carpel-sealc combined, the latl«r of tlie 
same constitution as that of Knes and Spruces, but perfec ' 
consolidiited an*! eonfluent with tie bract-scale.* 

513. In Cjeailscen, the tyiK of the flower of Aiigius|>erms is 
almost or quite lost ; yet the organs may i>e horoologizeil witii 
thtwe of Conifera.', which these plants are wholly unlike in habit. 

■ This liilcrnnl and ovuJitprous scale mny twm to br wbollj' h^pollicli* 
cal, uid u«uitiiiJ 1(1 lionioio^zc th« cuprtrtdineoUB wilh thf aUrlinratu 
cone. Without It, we ilionld luvc tn conaidor tlmt, whilo In AlHninnn (lie 
ovulci belong to Ivavca of a tecoaiarj nxis. in t'ii|irM>iii(!« ilii>]' niv Ihii-iii' 
nn ihiiio of > prim«ry nxis, or clw srp Mxillnr)- prwlurlium witliuut cKrjiclf. 
But b llic Amuoarl* Iribv tUv ititenisl ■rale it nbvious ; and there arc luffl- 

Fia. imsm. Zamlk. rlilrfly K. niHllk. Mtttj Itklianl, SM A inillo I>Uot. I^ 
LmrBrputoriRitlfl catkin. B71. A •lamm reninvsl. «hnwFngtinrnrrnn»»m»l1|«ll«i- 
nwauwlH (lio poIEala lup, sn. A h'niiila mkln. with a •luartcr (DL'tiiin cut mty, 
BJB A ftmiala dmrcr « carjBl, wllh two snlarilng otuIh nr imnng iw><l>, K*. Kltw 
•Md, wltli Itia thkk Iloalij' coat cut a*a; al ai>«K. SIS- LonsltuiUiia] tttUoa al tip* 


Their likeness to Palms and other Monocotjledons is confined to 
tlie i>ort of tlieir unbninchetl tiiinks and theiv pinnate leaves 
with iwrallel- veined or eiiiiple-veinetl leaflets ; nor have they any 
further resemblance to Fems, except tliat in some the leaflets are 
circtnate in vernation. Although a ti-opical type (of small present 
importance, compared with the part which it played in the 
Devonian and Cretaceous perioils), it has one small representa- 
tive (Zamia media, the Coontie) at the south-eastern extremity of 

the Unite<1 Stntcs, and a more striking one (Cycas revoliita, well 
known in cultivation) in the southern parts of Jnpnn. 

614. Following the analogy of Coniferip, each scale (whether 
of the pollen-bearing or the ovule- and seed-bearing ament) of 
Zamia (Fig. 569—575) is here regarded as a flower. Here the 
phylla, or scales with peltate top and stalk-like base, arc exter- 

cirnt inilicalions of similsr compOBition in the cuprcsulncons cont-BoIr* (o 
inducp iheadoption o( it by Parlslorc. who rejettcd the iilta of jrymnofpomi)' ; 
and. flnallv, (his I'nmpoaition il Bffinn<>d by StragburfnT (Diu C«Tiifi>rcn und 
die Gnetai-cen, 1S?2) upon microicopical study of the ilcvelopment 

FtO. BTIWrca. CurpnpliTUii of Cjran wvnlntm, mncli reiliic&l In iilro fi7(!. Ono 
b««Hn«rwoWbelnw»tMll™fle(iiorl™r-lnbo«t<«ran1.lhriir". 077. A Blmllarmrpo- 
phjU vitli InMntm mliired lo mere Icrlh, uul ovnln In plnco of tho lnwpi letth. 
STR. A dmlliir csrpophyll In matnn (tuctUluatlOD, beulns the Urge Jrnpaeemu nak«d 
totOM. Hm lut two after Blcbacd. 



sovrral at or on thr haw or rwh rono-nolr. alwaia wiiV "-I™ 
ii|manl. Arlmr-Vitw (Fijr. Silfi. .'»«;) Iwum .iwitU- |pajf ■ f i .« 
t« tin- m-ali': JimiiN-n', miiiM'iinuti urily itm-: inn- 1 »|.r.-w. w 
in Fiji. .'iiW),<>ni'iiiiil"Zi'iii>rnM>n'. At fliiwvniiiiiirii'. i.v •-*- 
HcnIcH iiKwtly njUK-nr an if itiiii|)lc: Uil in iwM ;i>-i>tT> t;,»T «.« 
tliifkcn (in-utlv wiiliin: nml tlicy nn- uoiiall\ iiii.lrr»!.-»: • » 
fomiNi-uil iir )>rnrl nml i-aqH-l-M'alf ii)i»>iiiMil. tli<- Ut:*r • f -;« 
saiiii' (iiiiHliliitiiui (It tliul i>f I'iiifit atKl Sjinnvn. 1-ut |»r'.-r.i 
i-oHHdlkliitttl tmil rtinfliwnt with (he brwl-WBli'.' 


'iin. In Cfradwer. tin- tyix- nf tin- (I<iir«T of Aiiiii— |- nii 

iiliinut i>r <|iiilf l<i»t ; vrt tin- nr^iint nuiv Ik- 1 lol-v"") ' 

tlio^.' or('«iiif>-r;i-. wlii.'h tli<-^> iii.iiit^ tin- nIi«1U iinlilti' in >o 

riii< itil.riinliiti-t .>viilifrn.i» tr»\v niMv m-. m li< l.i-i>h.illt ht;.'^;. 
ml :i-iiiii.<] l.> 1i»rii.>]<iiHzf llic i'ul<n •.tn.'.,u> oilli lh> '■>■'.. ;,t.. ^. 
WidK.iit il, »,- .1iu<i|.| Imrr i.i .ini.i.l.x llial. «li<lr in Atf tit-.-w ■•■ 
.U1..I1L' i..l.:iv....f a ..i-..n.l.rv Bvi.. ill C.i|.r, ..ii).»il» , ■» I. t. 
.., .,( :■ ,,ri,i,»ry «,.-, ..r .-l^- »n. »iIUr< ,^.-1,1. ti.«..».ui . .r,. .. 

-••I. null ih.' iiiuv II 

t- -1. in-r* rliliUKnl 

fTsiaicm: .ir r;i.->.- 

^T-Ei-iev - ' 

TWz- TOnnwnt TWn*- »t ■iTits Tli-.n 

,..,-,_,■, (R. p. ^-Hii:!,^; I, 

the pre: iiT a»?r nn'mui-ii-; n^ii.- : 

^1.. 1^1-.- i.:ma.!- i-a " :■* 

with T«mlfe-,- tawK -r -^aji-B— -:ii-. . 

iJc-i. I. r :a-- -Jr^- j.r; 

Itartaer t««to'oi«^ i. 5-.^^. -.ts--.- ::. 

:-■ :i * iin- \lr. fU^-l- lit 

cinr^BSk n v-c&cn a. A.nA'>:ir: : t*" > 

I. — 'r 1 saa^ i(»;-*>-'ir 

i^vcuam. ?iiiu<B>r>c v~:i: i^r tot: 

^■U\ 1 r i.a ■ -•. :i :ut 

DtTi.-aiita.SBc: : >'=i»----«i- •*-...> :■ i 

ufr ■.» ^as^ 'rr-(^-^'\a^ 

tin Zunut ibhLl Ue .' xuir b: t:r ~ 

^\T.i-K-ii<\:n -.Z.I'HIIll" nf 



' * * 


-^^i ,-^- •'- 

^>. / \ 

f n 




/ V 


SI .— 


tat 7iiii.*[ *■=!-*. mil i m--?" ^rrii-jr 

nf '—J.- ''■■- iiirj. ^- '. 

kanv-i □ -ii,-rT:a:tn a i:r - .::-^.--n -i 

a-".' ■' .i.-mi. 

::» J i'lTTit Ur r.j- . .-■ .-■ . :ii: 

:— s -.1 : --..r T-1.-.T:. - 

rf -^ y.-:^-^.^:^ -r i:r .-^r- fli.: 

Zunut 1 1; .v,5— 't: - ;,.-- — i:ar.i- 

i.- i t- <n- ii- " ::- 

l«i;la. .r -a.«> «^^ j-niiv -. ■! i:.. - 

^:i-::i.- 't^- i.-- -"■^- 



imllj' rouoh alike in the two sexea, which Ihrougliout the f 
octuiiv B«'i»irat« plaots. Tiie male flower (Fig, 368) or a 
ir it may l>e bo termed, bears indefiiiite pollen-sai» on t 
§i<Je or the peltate portion, sometimes extending to tl 
pari of its atallv. The homologous female flower, or earj 
lieurs a suspended ovule on eaeh aide of the stalk (Fig;. 573), 
whieh becomes a large fleshy-coated seed. In Cycaa the male 
anient is not very diitaimilar, althongh on a larger scale. 1' :' 
the i-flr[K)pliyll8 are evident leaves, not condenseil into &n «mi 
but loose or spreailing, of a character and aMjiect intcrtuc^l:. 
I>etween the lax bnd-seales which precede and the pinnat<.> folium 
l.^uves which follow them in development. Along the Da!i!_ 
of what wonld be leaf-blade they bear ovules in place ol" lent]. : 
lobes, or teeth (Fig. 576-578) ; and these, wbou fertilized tV.i i 
the male dowers, mature into lar^e and drupaceous naked Beeds. 
Even without fertilization, such seeds grow to their full size on 
the female plant of the common Cycas (or fulwly so-caUed Stiga • 
I'ulm), hut fonn no L'mbr3-o. ^^m 

Sectios VUl. Tiie Ovtle.' ^M 

515. Oniles (302) are peculiar outgrowths or production a of 
carpels wliieh, niKin the formation of an embryo within, becoRic 
seeds. In the angios[iermous gjTiQ?cium (4715) they are nor- 
mally produced along the margins, or some part of the margins, 
of tiie eariMlUry leaf (478), either immediately, or by the in- 
termediation of a placenta (485), which is a more or lees evident 
development of the leaf-margins for the sugiport of the ovules. 
Karely, jet in n considerable number of cases (iiOl, 302), ovules 
are developed from the whole internal suiface of tho ovary, i^r 
IVoni various parts of it, in no definite onler. directly from tin 
walls, and witlioiit the intcnention of any Unug whieh can h, 
regarded aa placenta. In Gj-mnoa|ierm8 (501-514) the ovub < 
UK bunift on llie face of the earpellary scale or at its base ; or 
on teaf-margina, aa in Cycas : or, when there is no repreaenta- 
tivc of the carpel, on the cauline axis, seemingly aa a direct 
growth of it. {Mi*, note.) 

SIC. As tj> altachment, ovules are either sessile, »'. e. stalk- 
losfl. or on a stalk of theb' own (Fig. 582, 584), tlie Fitsicci.t i 
or Pooosi'KRW. As to number they arc either solitary, few, oc 

■n (fgBl. periiBpi fl: 



indefinite!}' numerous. They maj' also be indefinite or variable 
in number when not particulariy numerous. 

517. As to situation and direction within the ovar}', the terms 
are somewhat special. Ovules are erect^ when they rise from 
the very bottom of the cell, as 

in Fig. 580 ; ascending^ wlien 
attached above its bottom and 
directed upward, as in Fig. 579 ; 
horizontal^ when borne on one or 
more sides of the cell and not 
directed either upward or down- 
ward, as in Fig. 314, 315, 530; 
pendulous^ when more or less hanging or declining from the side 
of the cell ; suspended^ when hanging from the apex of the cell, 
as in Fig. 581. 

518. The body and only essential part of an ovule is its 
Nucleus. This in most cases is invested by one or two proper 
coats. The coats are sacs with a narrow orifice, the Foramen. 
In the seed, the closed vestige of this orifice is termed the 
Micropyle ; wherefore this name is sometimes applied to it in 
the ovule likewise. When the ovule has two coats, the foramen 
of the outer one is called Kxostome, of the inner Endostume ; 
literally tlie outer and the inner 
orifice. The coats themselves have 
been named Primine and Secun- 
DiNE, but with an ambiguity in the 
application which renders these 
names unadvisable : for in their 
formation the coats apiiear later 
tlian the nucleus, the inner coat 
earlier than the outer; and the 
name of primine has by some 
writers been applied to the earlier 
formed, by others to the external coat. The proper base of the 
ovule, from which the coats originate and where these and the 
nucleus are confluent, is the Cualaza. The attachment of the 
ovule to its funiculus or support, which in the seed becomes the 

e - 

no. 5TD. Ovary of a Buttercup, <Ilvidc<l Icngthwino, to »ll«ir»lay its aBcendlnj? ovulo. 
660. Same of Buckwheat, with an erect otuIc. C81. Same of Anemone, with a sua- 
]»cn4lcd ovule. 

FIG. 582. Diagrammatic section of a typical or ortliotropous ovule (nuch as that of 
Tig. 582«), showing the outer coat, a, the Inner. 6, the nuclou!«, c, the chnbusa, or place 
of Junction of these parts, d. (The coats are never so scnamted and the nucleus so ro- 
duccl in size as is represented in t!ii« mere diagram.) r>8n. An ovule similar to the 
preceding, but curved, or campylotroiious. 6U. An ampldtropous ovule. 


TEZ n/>^rxB. 

llrL'if rr v.-ar. :ii*s i>:- :iU iJ^=»r: i- 'Jir ot-iI'^. In ibe simplest 

T',r:. 'A o- „- i* ii. Y\z. >i'. ''"" - --1^:- s^ 1 : '' / . a/a ar^ one. 
^y^ aL-/ 1:. ^^--^ wL-r.-v :>: "•-•:; •':-•. .-. ilr ir.i'jnt-j, a.* :n Fig. 
-S--;. I; ,: v.:r.' 'x-riiiU'. r^l"' ".!•■ ' li "^- ' i::i .Lz>:-:- »L:oL :<-^-ijme3 

xxi\ .>7. T^r.-f: ■:.•: Lll-::: *> liT'.ril. ' -: :--;■ :ji'.A2i a: :ho Lir^r 
<-n i. *..- tw'j "-iri:^ '■».r..vr';:r-i ' v a -i -r: rl '^r : ii.i in Fig. 088 
ti.«- "*'i ar»; i!<-^»iiritr«l ?•} thv wij«.-li- ivn^r^i * f t;.v v\"u.Ie. 

./K«. Tm: •irx.j/.— t ari'i :::•.•-: n ':il -:.:^r;.' ovu'.e :* :hat with- 
out a '-'^at. a^ in Mi*t>t*x- airi :Lt- «L- ".v i-rirr Lrriu'.hatx-iV. aiHl 
ill **ar/a!a'.-*:a.- ar*'! riri*-;a'.-»:n:-. I'lr.^ iia> f«:-rn ealli-»i a iiakeil 
ovular: hut I'^r»:r N-f-.-r*.- ovuW* of >urL '*;!iit licky weiv known 
thi-s Urnii ha'l li«-<-n aj^pn-jirlat*:^! to th-sr vf ^TX-miK^^i^enn-a. in 
th'r M:rii!<- of d<-^titut<: of ovarlal or iH-ri'-aq-ial o'Vi-riiig. i". ^. to 
\iu*:*i\i:Ti-*\ orttU. not to uiK.-'jven-'l *!»*<*•»/*. Tho ovule «:x>nsist- 
irijf only of nu«;i<-ij* may }«• t«'rnK-«.l carter Alph. Dt*C'aiidoUe) 
MunjfU, or \>i-\U'T at-ldmn^^om} 

'rjh, Th«' tiiiiicat(<l or chlainyili-<:nis rtviil.- is of tlirt-o priiii.*ii>ai 
kin'!-, with ono or twu suUjnlinato ni«iiifii.-aiion>. Those are 
Wnt orf/iofropo*/Ji. mmpyJot nrpoui, aii'l nn*ifn»p*iu$, aiiil the niodi- 
fif-ation *jil!«''l half-anatroi^ius. or amphifr^ponsr 

:f'J\, OrthotropooH rFijr. .>0. :>f<-2. ."i*^.";). or strai;jr]il ovule, is 
lh<; sirn|;l<r;»t hul leajst coimiion siH.-i.k->. U-ing that in which the 


.M m; ^ 

rlin!.'i/.a is at th<' cvi^h-nt haso. and the ori fu-i' at tlio <>p]K)site 
i-xlri'iiilly, the wholi* oviilr htrai<;ht (as the first i»ait <>f tlic naino 
(li-notfs) antl hvninH'trical. Atmptnts^ nu-aninir not tiirnod at 
ait. iH a latrr an<l ctvmolotricaliv niurli lu'ttcr nanii*. Imt it has 

1 All I'pidfTriinl Ktratiiiii ur tci^iinicnt niiiy not 1m' waiitinir to siifh nviiloji, 
foriiiinu II hort of udhcn-nt c-owriii^; l)Ut \\\U in nature and ori<:in Ik not 
Hlinilar to tlu- ovular coatM. 

'^ In I Jitiii foriii. oilhiifrnjHt^ mwjfiflnfinini, itmitrofHi, lunfJiitrnjHt^ — naiiH'< ij'ivon 
hy Mir)H>l, and ri'fcrrinK to tin* way in whicli tlic ovul«> i> tun,*i{ ritluT on 
ItNcif or on itN ftiipport. Sonu* Phij^lish botanists incongruously write wtho- 
trofttii, ninifitfhilrojMii, &e. 

FIO. BKn. i)rtliotr<i|M>iiii or Atri)|wiiiii oviih' nf Itiii'kwli«'rtr TiXi;. ('':iiiiiiv1iifn>i><'knii 
OTiilii iifrhlrkwiHHl. r*M7. AmptiltroTNniii oviilo nf Msillow r»ss Airitrin^xis iiviilonf a 
VIoli't. 11i<< iHliT A iiiilli*ut4'M tlir lilliiiii; r. tlif r1i:il:i/H. wliirli in 5»0 and t>>!^ curre- 
»|Miiiilii to llui lillum;/, tbo foramen or orlflcv; r, the rhapliv. 

OVULES. 279 

not come into general use. This o\'ule is characteristic of 
Pol3'gonaceaB, the proi)er Urticacea?, Cistaceoe, &c. 

522. Campylotropous (Fig. 583, 58G) is the name of the ovule 
which in the course of its growth is cur\'cd on itself so as to 
bring the orifice or true apex down close to the base, here both 
chalaza and hihim. This and tlie orthotroix)U8 ovule beg^n 
their development on the placenta in the same wa}-, but the 
campylotroiK)us develops unequally, one side enlarging much 
more than the other, esp3ciall3' at the base, until the ovule 
becomes reniform, and chalaza and orifice are brought into 
close proximity. Canipylotropous ovules are characteristic of 
Crucifera?, Capparidacese, Resedaceai, Caiyophyllaceie, and 
C henopodiacea* . 

523. Amphitropoufl (Fig. 584, 587), also termed Heterotropous 
and sometimes Ilalf-anatropous^ is between the preceding and 
the following ; and it passes in various instances either into the 
one or into the other. The body of the ovule is straight or 
straightish, but it stands as it were transversely or at right 
angles to the funiculus and hilum ; and it is fixed by the middle, 
the chalaza at one end, the orifice at the other. An apparent 
continuation of the funiculus, adherent to the outer coat, extends 
from the hilum to the chalaza. Compared with the preceding 
form, the explanation is, that the unequal development at its 
formation is confined to the basal half, and the axis remains 
straight, while the whole is half inverted by the ver)' unequal 
growth. Compared with the next form, the inversion is less 
and the later growth or extension of the apical portion greater. 
The amphitropous ovule is characteristic of Primulaceae, and 
is common in Leguminosse. 

524. AnatropoDS (Fig. 588, also 579, 581, 597) is the name 
of far the commonest siKJcies of ovule, that in which the organ, 
under the course of its growth, is quite inverte<l on its base ; so 
that, instead of standing at right angles with the funiculus, it is 
parallel with it, or rather with the apparent contiiuiation of 
it, which is adlierent to its surface as a sort of ridge or cord 
extending along the whole length of the ovule, from hilum to 
chakza. The latter occupies the seeming a|>ex of the seed ; 
and the oi^anic apex or orifice is at the other end, close beside 
the hilum. At maturity, the ovule is straight, but not wholly 
symmetrical, the attachment being oblique or somewhat lateral, 
and the ridge or conl on that side not rarely prominent. 

525. The cord or ridge, which extends along the whole length 
of the anatroixuis ovule, and for half its length in the amphi- 
tropous (Fig. 588, 587, r), is named the Ruapue. This is not 

giUt andiM 
>, or uC flOoP 


at all n N.-aiu. an tho Greek word denotca. lis origin^ a 
wliuk- Kli'iictiliv (if iiiH!li o\ulea will be apprehcuJwi liy t 
viirluu* Bt4igi'B of it« givwlb. 

62G. All ovule of any kind at the bcgmi 
(■xprf8C!t!noo ov outgrowtli of tiit plauutila, o 
pari of tbt' Ifiif-surface if [liere is no (levcIo(icd 
lilniimta. Tills indpieiit uvule ia the nucleus (514), 
f Hi lie nueletlH surmounting n nnliinentai;j- AiintnilUft. 
'I'lu- nuelona ia aofl cellular tissue ouly, fWrni tifst U> 
/A j IjiHi. Tlio aehlamydeous ovule (319) umUT|Eo*« no 
J I tm-iiwT ^ll:^■l•I■^Ill^pnt except in size or shajw. Indeed 
Wy Borni'tiini'S (ns in Balanophoreie) tliis bttre aucleus 
In rwluced to a few cells of pai'encli)-ma. 
AS". In ordinary ovules a nevr growth early begins arotind 
the liflMt iif llie nuclciiM, or is Bometiuips toKaneotis wilb it. at 
Ihitl an Ik iii»;{ (or part of a ring), soon as a eup, at li-ngtti as 
an oni-loMlnit tine or eovering, open at the top : tliia is the inner 
t'oat of lliti nvult' when there are two. The outer coat begins and 
((iH-* on in tim name way. unU »t length grows over and ent-losra 
lliu InniT coal ns Lbat diil the nucleus. (Fig. 590-59ii.) When- 
ever tliert" ia a third and more exterior eoat it is forine<l iltiring 
tlui growlli of tlio fertilized ovule into the seed, to whieb tliere- 
f(in'lt killings, and in which it takes tlic name of arilliia. (o'JT.) 
At the lime of fortillKatlon the ai»ex of the nucleus, or a i>r(>- 
longatliin of it. unnnlly prujet'ts lieyiiiid the orifice and there 
reoolvcH Ihc dineetidirig pollcn-tulw. Some fibro-vascnlar tissue. 
r»iM'cinlly spiral ducts, may be found in the funiculus ami cha- 
lu^ta. someUmea csttcnding into the coats. 

(I'iH. Tlie develapmeul of tlie ortbotrogxins or atropoua (iin- 
turtied) ovule prot^etKls symmetrically, witliout distortion, tlie 


part* keeping their primitive dirt-ction. In the eauipytoiro]ion-- 
Ihe whole of onu side of llic ovule greatly oiilgmwa the otlnr. 

aboat a 
only niu 

tWU. HiignlB»l V 
DionUi loifhru M 


■r »» 

> Aw 'la 

>n -t a Mrfrl or MngnMlia I'mltnll*. 
fil>sii>oiiu.-rhi »vn]«.aC(l.tal]ni« 

il" nr MBpinllB Vnibn-n.. .l.™inB iIm 
Ovuk B Hi-ik nlilH than In IMi Mil. 
> lain an Same tr-m a nwrtr f"l"- 

tfataainii |S«*<I(»r. IMM 


BainD t 


>n »r Ui4 «wU iMd il>» atiilm 
i*«Ie nr t«.i l«l*r. 001. 8am> 
"war-l.iiil. IM. 3<wi)iattln*nr 
l»ufU.<irlwt<)>«- m. Ciw-d 

OVULES. 281 

In the anatropous, the inequality of growth is mainly confined 
to the base or chalazal rt^gion, which ends b}' becoming upper- 
most ; and the full-grown ovule has the &\> 
pcarancc of being inverted on and adhercnt 
to the upi)er ^xMlion of its funiculus, the 
rha[)hc. Fig. 5^9-59 7 illustmte the coui*se ^ 
of development from a comparatively early 

529. The direction of anatropy or of other 
turning of the ovule in the course of growth 
is somewhat* diverse. But in general, when- 
ever ovules are in pairs, the two turn from ^ 
each other, in the manner of Fig. 315, and so present their 
rhaphes back to back. The rhaphe-bearing may therefore be 
called the dorsal side of the anatropous ovule. The same is true 
in the case of numerous ovules, viz., those of one half of the 
placenta (or one leaf-margin) turn their backs to those of the 
other. When such ovules arc solitary or in single rows, and 
either asc*ending or hanging, the rhaphe is usuall}' on the side 
next to the placenta or ventral suture, as in Fig. 57G : it is then 
said to be ventral (i. 6., next the ventral suture), or adverse to 
the placenta. In certain cases, mostlj* in hanging ovules, as in 
Fig. 57H, the rhaphe looks in the opposite direction, toward the 
dorsal suture or midrib of a simple ovary : it is then said to be 
dorsal or averse from the placenta.^ 

* By comparison of Fig. 578 with 576 and the like, it may be perceived 
that the difference is explicable by a kind of resupination of the ovule of 
the former. That of Ranunculus, if inserted higher, would become hori- 
zontal ; and if the insertion were transferred to the very summit of the cell, 
it would be suspended and the rhaphe averse, as in Fig. 578. Upon this 
conception, Euphorbia and its allies has normally suspended ovules, the 
rhaphe lK.'ing next the placental axis, and Buxus and its allies, resupinately 
suspended ovules, the rhaphe averse. The propriety of regarding the ad- 
verse rhaphe as the normal condition is confirmed by the fact that the only 
instance we know of solitary erect ovules from the base of the cell having 
the rhaphe averse is that of Uhamnus and its allies ; and here it was shown 
by Bennett (in 1*1. Javan. Bar. 131), and confirmed by the analyses of 
Sprague (Gray. Gen. 111. ii. 108, plates 163-100), that the rhaphe of the young 
ovules is ventral, so that the dorsal position, when it occurs, is the result 
of torsion. J. G. Agardh (in his Theor. Syst. PI. 178, &c.) maintains the 
contrary, l)ut is not sustained by later observers. 

Accordingly, even if we adopted Agardh's estimate of the botanical 
value of the characters here considered, we should prefer to express these 
differences in the phraseology above indicated, and not to adopt his terms, 

FIG. m:. Same aji 5iK more m»inilfte<1 ; the outer coaf (a), the Inner (6), nucleus (r), 
and the bundle of spiral ducts {d) in the rhaphe (running ft'om placenta to chalaza) 


580. Origin and Nstore of Uie Ovale. It bns been i 

stated in gc'iiofal terras llint ovules &ve peculiar outgrowths 
or |ii'0(UictiuiiH, generally of tlii? uiui^iis of oaqieUury leaves 
(515) ; tbat iLL'y are eom[H>8e«l ol' pai'eiicliytnatous cellular sub- 
stance, at least as to the iiuekus, of wliicli the simplest ovute 
wholly consists (52G) ; that the coats originate sulisetjuently to 
the nucleus ; aud that llie outer coat is ul' later origin than tlie 
inner one. (.ilB.) The mamililhrm protuK'nince of wliii^h the 
forming ovule at first consists originates iu one or uiorc cells uf 
a layer directly beneath tlie epidermiB.' 

531. The moq)hological nature of the ovule has befn much 
discussed. The commonly prevalent view was that the o\ulo 
is homologous with a leaf-bud, and that its natui-e is in some 
degree illustrated by such buds as those whieb <le\'elop on the 
margins of Ihe leaves of Bryopbjlhim, as shown in Fig. 322. 
But such buds, and the bulblcts or fleshy buds which a|)|>car on 
the fiiee of certain leaves, follow the universal order of budding 
gntwtli, tbat is, are ceutri|)etal in development, the outciTnost 
[larla Inking the earlier and the inmost the later forrae<l. The 
ovule, on the contrni^', is basiiietal or centrifugal in develop- 
ment, the nucleus being Brst aud the outer coat Inst formed ; 
tliercforc the coats arc not homologous with sheathing leaves, 
nor the nucleus with a vegetative axis. The ohh-r theory lias 
accmdingly giien way to the present one, in which the ovule 
answers to the lobe of a leaf |M?eutiar!y Iransforuied, or to an 
outgrowth of a leaf, whether from its edges or surface. The 

apehvpoat, rpUnpaM, and hrtmbvpoat (Ihe drat Iwo new, the lut emplnyed 
in a new scntc). Uie more mi eincu the ftpplieatioti is confuaeil with hy[H> 
ihellcal connidoralinn* «nd the nrieetily of bringing tlie ovults ideitlly l»»i;k 
to ■•cending or hnrizoiual poiilioiu, ll majr be staled, briefly, IhBl Urirro- 
tro^u>, lu AKtnlii'a terminology, Rpplio* to the noniiBl position of i-ullnl- 
«r«l ovatu, with rliaiilics iMuk (u liauk, in opposite dirMlions on the two 
halvn of ihc plnecnlR : Apempout, to an erm;! or ucendiDg ovule wiili iti 
Tliaphe next the plaL-enlal nxla. and a lianging one liaa its rhaphc arerve 
froiD it; EpiiroiKmi, when an prvct or ucemling ovule has its rhaplit! avene, 
■ad a hnDginjjE one lias it nilreric. 

> nofmebltrr'* ttatemcnt tliat the nimple ovule ai Orchis oriitiiMles in 
the division of a Anif\e rpldermat celt (and' ia tlieretoro a tridinmr) \t con- 
lrorert«d liy BtnuliurKtr and hy Warming. Tlie lalt«r adJi the nniarli, 
lliat even if il wi-rc >o in eases of pxircmc simptiHIy. itda trould not intall- 
dale the propottllnn that tlie ovule is to be rcmnled sa llie bomnlnjiue uf Ilia 
lohc flt a Iraf. Such a tobc is not rarely rcdurcd to a single bHiflle. For 
the whole inlijpoi of the origin, devclopineni, morphology, and tlieory of the 
oTiile, see Warming's very flaboratc and perspicuous meniuir. Dc POvule; 
alto (he paper* of CuUliowaky, Van Ticgbem, fie., rufttrml to in tiotn to 
paragraphs (iOO. W\. 

OVULES. 288 

great advantage of this viev is that it serves to horoologizc the 
(hictiflcation of flowering I'lants with that of the higher Flower- 
less rianla, or the Ferns, the sjxirangia or analogues of the 
ovule heing outgrowths of the leaf.' 

o3-2. Orlgrlnatloa of the Embrjo. The whole process of fer- 
tilization and the resulting produc- 
tion of the emhryo, also the history 
of the subject, belongs to the suc- 
ceeding vohimc, involving as they 
do questions of minute anatomj' and 
of phyaiologj-. But a general idea 
may here be given of the way in 
which the embryo originates. The 
tube whioh a grain of jwllen sends 
forth into the stigma (574, 57a) 
penetrates the style through loose 
conducting tissue charged nith 
nourishing liquid, reaches the cavity 
of the ovai'y, enters the orifice of an 
ovule to reach the apex of the nu- 
cleus, although the latter sometimes 
projects to meet the pollen-tube. 
Meanwhile a cavity (the tmbryo- 
tac, which is fonnc<l by the great 
enlai^omont of a single cell of the 
tissue, or of two or more cells the 
proihict of a mother cell) forma in 
the nucleus, the upper [Mirt of it 
commonly reaching nearly or quite 
to the a|K'X of the nucleuH, which 
the iH>llen-tu1>e impinges on or ^ 

sometimes i>cnetratcs. A particular portion of the protoplasm 
coulainitl in the emhryo-sae forms a globule, and this at the time 

' Till- nrlvcical('» <if ilili view naturatl; maintain that ovulm and placenta 
always Ik-Iork to ltav(.g, and never truly to a caulinc axis ; tiiat in tlie pro- 
central plaivntatinn <if Primulacce, the actual oTuliferoua surface is an out- 
gniwili of the Uueg of ilie carpcllary leaves cnaltwcenl with each other and 
adnalc to a prolongation of the torus ; also that in those Gymnosperms which 
have no i-arpophyll, such at Yew, the whole nascent carpt'llary loaf, or rather 
tlie papilla which would otherwise develop as such, is directly developed into 
ovule. This, beinj( solitary and the last production of the axis, necessarily 
appears to terminate it. IWXi, 001, notes.) 

with Inniritiulinal 

ii'IO.GM. DUcnm rpiavHTiIlnEBmsffnKlBl rli<<ll<<rBu 

one (tain illilinrllT ulimrlnii IIh tul>«. whii-h hut fMifimtM 
cailtr of (he ovary, cnleml Ibe mnalb of tba solltuj' i 





of fertilization Is {band at the n|>ex of the sac. at or lU^acent H 
the part reached by Uie polleii-tiihc. Not rort-ly it iidhorcs U> the 
j» on 051 ns OD wall of the sat- exacUj' 

op|H)sit« tlie termination 
of tlie )Mlli.'ii-tuhc. This 
13 caUed the embryonat 
reticle. To it tlie con- 
tents of the iKiIU-ii-tube 
are ill some manner trans- 
feiTiil, Upon which it 
lakes a more deRiiitc 
slia|)C, anjuiivs a wbU of 
ittUnlose, and so beeoiiies 
n vegetahlc t-ell. This 
di\ ides into two, the lowei- 
apiin into two. nnd so on, forming a L-hain (tlie lutpmsor or pro- 
embryo). The temilaal cell of this divides again and again in 
tliree directiona, prodni-ing a mass of cflls which shaiies itself 
•M ouB aa m into the einbrjo, the initial 

ad A /^"^/l I>lant of a new gcnera- 

/ 1 /J M r iVi^^ X'ion- Ordinarily the su»- 
|K!iisor soon disappeara. 
It is attached to ijje ra- 
dii'tihir end of the em- 
bryo, which consequently nlwnye jiointa to the foramen or 
micropylfi of the seetl. The jH-oecBa in GymnoBpctms is more 
coin[>lex. and has to bti separati'h- deBcribcd. 

533. Polyemhryony, the production of two or more emhrj'Os in 
one seed, is not uncommon in (lymuoBgwrms (tliere being a kind 
of provision for it), and is of uccueional but abnormal occurrtrnc'e 
in Angiosi>ei-ms. in the 8cc<t of Mistletoe. Sanlalum. Ac. In 
these it results fkoni tlie production and ferliltzation of more 
than on« embryonal vesicle. Straslmi-jrer has retently nsccr- 
tained that the commoner polyemhrvony m the seols of Onions, 
Oranges, Fimkta, &c.., results fhiin Uiu prudiiclion of adventlve 
cmbrjoB. wliieh originate in the niieleus outsiile of the embryo- 
sac nnd wholly inde{>endent of R-rlilization.' Two kinds of 

' Slraaliuriipr. t'cbrr I'olycnihr^rnnip, tn Zriuchr. Nnlurwii, Jcriit. yt\. 

1878 [kv Atiiw. Jcmr, Sci, April, 18711). Il wu lound tliat wlien, by i-xi-lo- 

■fini of pallcn. Ihr fnmiation of > ivimial t^mlirTn wm preTented, do ndTentivc 

Fin {M>. DtMTMn iif llie in»iimmr ui.l InrlpMit nnbrjw ■[ Ila utmnltr. AM. 

Tha MUSI, ullti tba •nhrya n Illl1« man •Involnpnl. ml, Th* mine, mnn <1inBln|,«<I 

iit<>Mylv(nM Ihlnllj lii>11«t«l U Ihe lover mi. im. Same, ttlUi llH |in4plHit 

MTilAst. n>3. TIhi Rlnlir^', nurlr nimHo'ol. 

w cutyleduut Full/ iI*v(i1d|>mL 

tttl. Suue, ■ 


anomalous reproduction are therefore now known, wliicli are 
intonnediate l)etween sexual and non-sexual, between budding 
and fniiting propagation, viz., — 

Apogamy^ which is budding growth or prolification in place of 
that which should subsen-e sexual reproduction. This was dis- 
covered in FeiTis by Prof. Farlow, while a pupil of De Bar}', by 
whom our knowledge of the process has recently been extended, 
and this name ira[)osed.^ The production of bulblets in place of 
seed or embryo answers to this in P'lowering plants. 

Parthenogeny^ the counterpart analogue of apogamy, is the 
non-sexual origination of an embryo extraneous to the embrj'onal 
vesicle or even the embryo-sac. However abnormal, its occur- 
rence is probably not so rare as has been supposed. 


THE FBurr. 

Section I. Its SxRucrimE, Transformations, and Dehiscence. 

034. The Fmlt consists of the matured pistil or g}-na?cium 
(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 ovar}-, to a cluster of such ovaries, at least 
when somewhat coherent, to a riixjned ovar}' with cal^'x and 
otlier floral parts adnate to it, and even to a ripene<l inflores- 
cence wlien the parts are consolidated or compacted. Fniits, 
accorrlinjrly, are of various degrees of simplicit}' or complexity, 
and should l)e fii'st studied in the simpler fonns, namely, tliose 
whi<*h have resuUed from a single pistil. Such a fniit consists 
of Pericarp with whatever may be contained in it and incoqx)- 
rated with it. 

embryo appeared in tbom> seeds which habitually produce them. To this 
Cjrlebojryni' offers an exception. The female of this diwcious plant habit- 
ually matures fertile setnls, 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. Parthenwjfnesin, 
of which Caelebog>'ne was the most imequivocal 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.OB; De Bary,Bot.Zeit.xxxvi. 405-487. 


535. The Pericarp, or Seed-vettel, 19 the ripened ovary. It 
should, therefore, aeeoi'd in stnieture with the ovary Ctom which 
it is derived. Yet altoralions sometimes talce pUiee during fhic- 
tificatioD, either by the alKirtion or obliteration of parts, or liy 
accessor}' growth. 

536, Internal AltcraUons. Thus, the ovarj- of the Oak con- 
sisU of three cells, with a pair of oviilea in each ; but the fhiit 
has a single cell, filled with a solitary seed, only one ovule being 
matured, while two cells and five ovules are suppressed, the 
remains of which may be deteeled in the aconi. The ovaiy 
of the Chestnut has six or seven cells, and a jiair of suspended 
ovules in each ; but only one of the dozen or fourteen o\-ules 
ever develoia into a seed, except ns a rare monstrosity. The 
three-celled ovarj' of llie Hurseehestnut and Buckeye is ^niilar 
in stmcture (Fig. 608-611), and seldom 
ripens more than one or two seeds ; but the 

abortive seeds and cells are obvious in the ripe (Vuit. The 
ovary of tlic BiR'h and of the Elm is two-celled, with a single 
ovule in each cell : the fruit is one-celled, wttli a soUtary seed ; 
one uf tile 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 ovarj- are not uncommon. 

037. On tile other hand, there may be more cells in the ft'iiit 
than ttiere are primarily in the ovary. Thus the fVuit of Datura 
is dicar])ellar}- and normally two-celled, with a large placenta 
projecting from the axis far into the cells. But each cell be- 
comes bilucfl/nle, tliat is. divided Into two, by n fUlso partition 
growing out fVom the back of each carpel and cohering with the 
middle of the adjacent placenta. 80 tlie 5-carpelIary and nor- 
mally fivc-i-eUe*! ovary of common Flax early becomes spuriously 
ten-ccll«l (morphologirall.v speaking, not lO-locular. but 10- 
hcvUale), by a false partition extending from the back uf each 

ir wrurj ntn Bnckrye I*kiiIim Pmin), >linwtin[ 

no. TruidSrwHKtl.nnf th<^MRia<Ui>|.liiyln(iU 

n. Sunt of Imir-growti fralt. wl'h •Incln frr'ilo ■»!, •!«>[- 


carpel across its cell (Fig. 539-541) ; and the solitary caqwl 
is similarly divided lengthwise in many si)ecies of Astragalus, 
as in Fig. 534. Transverse divisions or constrictions across 
a maturing ovar}' (such as is seen in Fig. G20) are not uncom- 
mon, esix^cially 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-G27) , 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 BoiTaginacea?) , 
which thus l)ecome burs and are disseminated by adhering to the 
hairy coat of cattle. All these are of the nature of sui>erficial 
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, necessaril}' makes a constit- 
uent part of the fruit, in the pome (575) doubtless a verj^ 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 Compositae for dissemina- 
tion. Or, in small pericarps, the style ma}' persist as part of the 
fruit, and subsen-e the same ends, either by becoming featherj' 
for aerial dissemination, as in Clematis and in one section of 
Geum, or b}* becoming hooked at tlie tip for adhesion to fleece, 
&c., as in other si)ecies of the latter genus. Or adjacent parts 
which are not actually incoqxjrated witli the pericarp may play 
similar parts in the economy, as the hooks on the calyx-tube of 
the dry calvx of Agrimonia, which at maturity is detached with 
the inchulod fmit, the flesh}' fructiferous cah'x of Gaultheria (Fig. 
C51) and of Mulberry (Fig. 654) ; and the pulpy fnictiferous re- 
ceptacle of the strawberry (Fig. 653) : the ultimate utilities in 
lx)th classes of instances l>cing similar, viz., wide dispersion of 
the seed by animals, whether by external carriage, or by being 
devoured and the voided seeds of flesh}' 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 
cari)els of Columbine, and Marsh Marigold (Caltha) , or the poil 
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- 


out bat soft witbin, as in n pe\io ; ov fleshy or hcrrj-lilse witUotit. 
but itKluratccl within, as in all stone- I'm its, sut-b as tlie cbeny 
aiid iMraeh. 

511. When ttio wnlLsora pericarp consist or two loj era of tli»- 
similar tt-xture (as in a peacli) tlie outer layer is called KxocAiir, 
the inner Emhkarp, these tenua moaning exterior and interior 
partA of a fniil. When the external layer is a comparatively thin 
Btratiini or film, it is Bometimes termed the Epicasp. When it ia 
fleshy or pulpy it is named Sakcocarp, When the endocarit witbtn 
a §arcofarp is bard and bony or crustaceoiis. forming a sUell or 
stone, this is termed a Pl-tames. When three concentiic layers are 
dislinguishuble in a i>ericarp, the mid<lle one is called Mesocari*. 

542, Fmita may he divide<l into two kinds, in refercnoc to 
their dischai^ing or retaining the containeil seeds. They are 
dthiierni when tbey oj^n regularly to this end ; indehiictnt when 
tbey remain closed. There is a somewhat intenneiiiate condi- 
tion, when they rupture or buret irregularly, as in Datura Metel, 
&C. Drj" pcricar|)s with single seeds are commonly indebisccnt ; 
those with several or many seeds mostly dehiscent. Seeds pro- 
vided witli a wing or coma or any analogous help to di8|»ersion 
are always in dehiscent |>ericarp8. I'ermanently Hesby peri- 
parjw are initebiscent. stone-fliiits as well as berries. But in 
some slone-IVuits {i. e., with indurated endocarp and fleshy 
exDcarp), such as those of Almond (I'ig. ii4U) and Hlckorj*, 
the baiTly fleshy exocarp or sarcocarp dries or haitleiis, instead 
of eoHening. as malurily is approached, and at length separates 
l^ni the putamen by dehiscence. 

3-13. Dcblscenee. the o|N^ning of a pericarp for the discharge 
of the contained seeds, is regular or irregular; or, l>ettcr, is 
normal and ahnarmiil. For most of the abnormal or non-typical 
modes are as determinate and uniform in occurrence as the typi- 
cal modes. A good English name for dehiscent pericarps tn 
general is that of Pod. 

544. Regular or normal dehiscence is that in which a pericarp 
Bplit« vertically, for its whole or a part of lis length, on lines 
which answer to sutures or junctions, that is, along bnes which 
corrcs]>ond to the margins or midribs of carpellary leaves, or to 
the Hues and surfaces (or commissures) of coalescence of eon> 
tigiious cnri^els. The pieces into which a pericarp is thus suif 
dered are termed Valves. 

54."i. The normal dehiscence of a carpel is hy fts inner, ven- 
tral, or ovulifcrous suture, that is. by the disjunction of Uie 
leaf-margins, as in Fig. G18, Its only other line of normal 
dehiscence is by the opi>ositc or dorsal suture, that ia. down 



the midrib. L^umea iiBuallj- dehisce by both sutures (as in 
Fig. 619), therefore into two valves. 

546. A deliiscent iwrican* formed of two or more carpels is 
calle<l a Capslxe. Tlie two leading tcTms descriptive of capsular 
detiisccnce were based upon the modes of opening of peiicaqw 
liaving as many cells as car^jels : they are the trpliddal, tbat is, 
as tiie term denotes, cutting through the tepta or dissepiments ; 
and the tocuiieidal, that is, cutting into the loculi or cells. 

547. Seplicidai, the dehiscence tlirough the dissepimentsf is 
the disjunction of a |>ericarp into its constituent carpels, these 
then usually themselves dehiscing down theur ventral suture, 

as in Fig. 012, illustrated by 
the diagram. Fig. CIS. Good 
examples are furnished by the 
llj-peiicum Familj' (the pistil 
illustrated in Fig. 536, 537), 
where the plnccntffi which 
comiK>sc the axis are carried 
awny on the edges of the par- 
titions or introflexed valves; 
also by Rhodo<lendron, Kal- 
mia, and the like, in which 
the placentae remain combined 
into a column in the axis (the 
CoLL'UELLA or column), ftam 
which the edges of the valves 
break away. 
548. The septicidal dis- 
^ junction of the carpels doep/ 

Onot of itself open the cells. 1 
Such separated car 
,13 one-seeded not rarely remain 

closed, as in Mallow, Ver- 
bena. &c. Or when dehiscent they maj' open both by the ventral 
and dorsal sutures ; t. e., the pericarj) may first divide into its 
constituent cari>els, and then each carpel break up into half 
caqK'ls. as in Euphorbia. 

549. Liirulicidal, the dehiscence into the loculamenti. loculi, 
or cells of the [)cricarp (shown in Fig. 614, and the diagram, 
615), is tliat in which each component carpel splits down its 


FIG. G1J. Septidil^lr<letil»entIrleuiiclUr7C*pialeorElo>]aVlr(lnlc 

GI4. I»cullc|.lii]lr ilelilKcnt McuptlUrr okpculs of w 
It the middle, su. UtacnunoriocalkkUldaldMeuca. 

Iili, dlThled tnuw- 



dot«al Biititre, aa in Iris, Hibiscus, (Enotlicra, &c. In this, tlie ' 
dissepiments remain intact, it' tticy breiik nwaj' from tlic contre 
tlien tliej- are bonie on tlio raiildle of Uie lalves, as in the figures 
almve cited, irthej' remain eolierentin the axis hut break uwa}' 
from the calves, tlie result is one form of what is called — 

550. Srptifragal dehiscence, i. e., a breaking away of the 
valves fVom the septa or partilions. as shown in Fig. GIC. Tliis 
leiiiesentB the locnLcidftl form of the ae|)tifragal mode, which is 
less common thnn that of the nccomi>anying iliagram. Fig. 617. 
Here the [lartitions alteniale 
with the valves ; that is, the 
dehiscence of the iTericarp is 
/ J I I ""^--..^^^•^ 1 of Ihe aepticiilal order, as 

Y ^,,,-''''^~\^ /I T I near aa may be, but the par- 

\ y \ \ y tilions do not split, wherefore 

\» _../ >v. ,.-/ the valves break away at the 
'"" "" common Junction. To this 

the term marffinirii/nl baa been applied. It occui-s in the 2-3- 
carpellary capsule of IpomKa (esiieeially in the common Morning 
Glory), in the 5-cBrpellary capsule of the North American aiiecics 
of Bergia ; likewise in Uie 2-ear|>ellar)* [kxI of Cmcifera; (Fig. 
623), with a difference that the placenta; from which the valves 
break away are here jjarietal and the partition is abnormal. 

651. The terms seplicidal and loculicidal apply equally in plan, 
though not with etymological correctness, to one-celled cajMiilea j 
witli either parietal (596) or fi-ee central (59D) plaeenlte. When j 
the dehiscence is of the septieidal tyjje and the placentalion jMiri- 
etal, the (half) placents are borne on the margin of the valves, i 
as in the Gentian family and the si>e<?ica of liyperienra with one- i 
celled capsule. When the plncenlic arc borne on the midiUe , 
of the valves, as in \'i(ilt'ts, the dehiscence is of the loculicicUi , 
type. In the case of fVoe central placenbe with no trace of 
partitions, (he chamcter of the dehiscence may usually he deter- 
ininod by the |X)sitloo of the styles or stigmas relatiie to the I 

552. Dehiscence may be (jnite normal although verj" partial. 
Its when confined to the apex of the capsule of Cerastium and 
of Primula, and even to the jjores under the radiate atignias 
of Poppy. 

558, Irrtfftiitvr or abnirrmnl dehiscence is surli ns has no res|»ct 
to the normal sutures; as where the dehiscence is transverse; 

'. RuiDoufaciiikiiunr 


either extencUng part wa}' round, as in the pod of Joffersoiiia, 
or completely round, so that the upper part falls off like an 
unhinged lid. This circumscissiie dehiscence occurs in many 
plants of widel}' different oitlers ; such, for example, as Purslane 
(Fig. 621), genuine Amamnths, Plantain, Pimpernel, and Hen- 
bane. In other cases, as in Antirrhinum (Snap-dragon) and 
its allies, the cells burst b}- iiTegular laceration at a definite 
|)oint, and discharge the seeds through the ragged ix?rforation ; 
or one or moi*e neat valvular orifices are formed on some parts 
of the wall, as in Campanula. 

Section II. The Kinds of Fruit. 

554. Fuurrs have been minutety classified and named ;^ but 
the terms in onlinarj* use are not very numerous. A rigorously 
exact and particular classification, discriminating between the 
fniits derived from simple and from compound pistils, or between 
those with and without an adnate cal^'x, is too recondite and 
technical, and sometimes too hypothetical, for practical pur- 
poses. It is neither convenient nor pliilosophical to give a 
substantive name to every modification of the same organ. P'or 
all ordinar}' puqx>ses, botli of morphological and systematic 
botany, it will sufilcc to characterize the principal kinds under 
the four classes of — 

Simple fruits, those which result from the ripening of a single 
pistil ; 

AijgregcAe^ those of a cluster of caq^els 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 

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 lie distinguished, upon difTerenccs of 
texture, into Dry Fruit*^ Stone Fruits^ and Baccate Frvits ; or, 
better, into Dry and Fleshy ; and the first may be divided into 

1 The greater part of the forty-three eubetantive names of De«vaux'8, 
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. 476, and y. 0. 



cleliiaocnt and indehisccnt kinds.' Theoretically, cscb bdnd sisj^ 

be divided into those of a simple and those or a compound pistil, 
and some would make the primarj' division on this character. 
Some also would separat* fiiiita with adnate or superior calj-x 
fi-om those free of all such combination. But in practice these 
ditTcrences can seldom be indicated by snbsUiutative names. 
Tlie name of beiTy is equally applicable to the (Vuit resulting 
ft-om tlie single carpel of Aetiea, the synearpous ovarj- of the ] 
grape, and the similar ovary with adnatc calyx of a gooseberry 1 
and oninbciT]'. It slionld be understood that the kiuds shade I 
off one into anotlici- most fVeely. 

556. nehlsceut Froiti* (543), or Podt, are distinguishable into 1 
apoeitrpouf, or of single carpels, and syncarpont, of more than ( 
one carpel, i. e. the firat of n simple, the second of a compound 
pistil, The fltst kind is mainly represented by the FotHcle and 
the t^gitme; the seconit. by the Capmh and its modifications. 
A 557. A Follicle is a pod formed 

^W of n simple pistil, and dehiscent ))y 

^Sl one suture (this almost always 

BnA the ventral or inner sutm-e) alone ; 

^U na in the Larkspur. Coluiuliine, 
gr I'eony. and Marsli-Marigold (Fig. 
"" 618) ; also in Jlilkwcod and I>og- 

1>ane. There may be several follicles or only 
one to a flower, even in tlie same genus, as in 
Larkspurs. Cimicifiiga, &c. In Slagnolia 
(Fig. G48-G50), fleshy oaqtels Iwcome follicles 
dehiscent by the dorsal suture. 

558. A Legame is the pod formed of a 
simple pistil which is dehiscent by botti sut- 
ures (as in the Pea, Fig. Gt9), so dividing 
into two pieces or i-n/ce*. (544.) This is the 
fruit of the Pulse Family, accortlingly named 
I^iiminosie (Legiuninous plants); indeed, 
Uia name of bgnmo is restricted to the fruits of this family, 
and in desciiptive botany is extended to all the modiflcations 

' Dr. Moslcn'i mcxliflcBtlnn of Dicluan'i and HcMab'a claisiBuitian of 
•implc frulti, ft* Id primniy kind*, i* into 

1. jVu/1, or /l/imnoir/w, ilry snd indt-ljiiceni ; 
4. P'>dt, or Ilrnmamrpt, ilry. ilohiaccnl ; 

3. Slear-fmiit. or Pymmcurji. flMliy without, inilumied wllliin, iodeliiaoeat; 

4. Btrr'in, or Sttrromrju, flnhy tbroughout, Inilchiaccnl. 

?tO •"■ »„ 

D. LiniientufBl>«DUitlanv 


which that order presents. Some of these, in fact, arc in- 
dehiscent and reduced to akencs ; some break up at maturity 
into one-seeded indehiscent articulations or joints, which are 
dis|>ei-sed as if they were so many seeds. A legume of the latter 
kind takes tlie special name of Lohekt, Lat. Lomentum. (Fig. 
620.) In Alimosa (Sensitive-plant, &c.), such articulations de- 
hisce into two valves. They also fall away from the sutures, 
or from a persistent mai^nal border of them, or in some cases 
tlic valves thus Eull away entire. The persistent frame which 
remains has tteen called a Refldh, an architectural word, here 
taken in the sense of door-case. 

559. A Capsule is the pod, or dehiscent firuit, of any compound 
pistil. When regularly and com- 
pletely dehiscent, as already stated 
(544), the pod splits lengthwise into 
pieces or caliet. The modes of r^ular 
dcliisccnce are illus- 
trat«l in Fig. G12- 
017. Two modifica- 
tions of the capsule 
have received distinc- 
tivc names which are 
in oummon use, viz. 
the Pi/jnt and the 

560. A Prxls or Prxidlnm is a dry fhiit which opens by a 
circular line, cutting off the upper part as a lid ; t. «., the debiscence 
is<-iVcu/H«iM»7e.(5o3.Fig. 621.) In the Purslane, j 
Fimi>emel, Henbane, and Plantain, the pyxis ia I 
a capsule; in Amaranths (Fig. 637) it is a 
utricle ; in JefTersonia (Fig. 622) it is a modi- 
fication of the follicle, being of one carpel which 
dehisces transversely, and not all round, so that 
the liil remains attached. *■■* 

561. A Slliqite is a narrow two-valved capsnlc, with two pari- 
etal placentie, from which the valves separate in dehiscence : as 
in plants of the Cruciferous or iluBtard family (Fig, 623). 
to the fniit of which this term fs restricted. Usually, a false 
partition is stretched across between the two placenta, render- 

FIG. S2I. Pyxli of Panlue, FortuUt* oletMe* tLs lop wpukUng enllrelv uiil 
folllns nwny. 

Ptd. «a. PrxU-ltlie folllcniiir fruit of JvlTennnlii lUphjUk; the lid T«niiliiln( 

son. ^" 

ing the pod two-oelled in nii anomalous manner. A Siucle 
(SiVicu/a, iliminuti\-e of tilifua) is meK\y a short siliijup, the 
length of which does not inoro than twice or thrice surpass the 
hrcaiUh ; such as that of Shepherd'a-I'urae (Fig. 624). nncl of 
Luiiaria, Canilytiirt, &c. 

hdi. Indehlseent Ory Frolls are almost always one-scwlod or 
very ffW-seeiUil. If nnmcroua, the sc^Xn thus phiccd would not 
1x3 disiiersed. The ordinary kinds are strictly one- 
swilwl, and ui cominun latigunge are often cou- 
foundetl with seeds. The ways in which siK-h IVnits 
are dis|)ersGd are various. In the followiii);; case, 
Uie adaptation of the pericaqi to dispei-siou \\y wind 
distiiiguishca the species of fhtit. 

6ti3. The Samara, sometimes calleil in English u 
Aify, ifl an indehisccnt oue-seedcd fruit pi-orideU 
with a wing. In tlie While Asli 
the wing is terminnl (Fig. 625) ; 
in other species the whole frtiit 
is wing-mai^ned ; in Birch and 
Elm (Fig. G26) the wing sur- 
rounds the body of the [jericuip ; 
and the Maple frnit is a double 
samara or pair ofsuch fruits, con- 
spicuously winged fW)m the a})ex. 
.)IH. Akene (Lat. Achenium) is a general name for all the 
one-se«Hled, ib}' aud hard, indehiscent an<l setil-like small fi-nits, 
such as are jtupnlarly taken for naked 
seeds. But that they are Inie pistils, 
or ovaries riiicned, is evident fhmi the 
style or stigma Ihcy bear, ur from Iho 
soar left hv its fall ; and a section 
brings to view the seed within, provi- 
■" "* deil with its own proper inl«giimems. 

The name has been ivatrictrd to the see<l-like fhiit* of siiiiplr 
pistils, sneh as those of the Buttercup (Fig. 628. G29), Ancmom>, 
Clematis, and Geuin. The style in some species of the bitter 
mnains on the IVuit as a long and fcnthcry tail, in others as n 
short and hookeil one. botti being agents of dissemination. Tlie 
grains of the strawlierry (Fit;, ^'^^) <"* also akenes. The name 
is rxtendeil to all one-celled sec<l-like fruits resulting ftvui a 

FtO. «£». S«m»r« or kty of W]i 
Kim ntnii* .ln>*rfnin» <n 

Flo. «w. AebMitoa of 


compound ovaty, and even when invested with an adnate calyx- 
tube. Of the latter is the fhiit of Coapositffi. (Fig. 6S0-G35.) 
Hei-c the tube of the calyx is incorpo- 
rated with the surface of the ovarj- ; 
and its limb or border, olfsolete in some 
cases (Fig. 630), in others appears 
as a crown or cup (Fig. 631), or set of 
teeth or of scales (Fig. C32, 633), or as 
a tuft of bristles or hairs (Fig. 634, 


C3a), &c., called the Pappus. In the Lettuce and Dandelion 
(Fig. 63i!i), the acbcnium is rourate, or bcBke<l, i. e. its siimniit 
is extended into a slender beak. An akene with adnate ealyx 
bas l>ecu terine<l a Ctpsela. 

HGii. The I'tricle is the same as the akene, only with a thin 
and bladdery loose pericarp, like that of Goosefoot. 
(Fig. 636.) This thin coat sometimes bursts irregu- 
larly, dischaiging the seed. In the true Amaranths, 
tlie utricle opens bj' a circular line, and the upjjer 
part falls as a lid, converting the fruit into a small 
pyjTi* (560), — a transition form. (Fig. 637.) 

J66. A Cn7«pslfl or drain differs fh>m 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 IVuita this 
is the kind most likely to be mistaken for a seed. "" 

ufi7, A Sot is a hard, one-celled and onc-scetled, indehiscent 
fruit, like an achcnium, but larger, and usually producetl from 
an ovarj' of two or more celb with one or more ovules in each, 
all but a single oiule and cell having disappeared during its 
growth (J36) ; as in the Iliizel, Beech, Oak (Fig. 638), Chest- 

ex. Of Ibe Duidell 

AmaruiUi, bj tmwrraw d«hlK«iica bMnmluf a pjxls. 

m GooMfoot. G37. 1 

THE FKurr. 

Dut, and the like. The nut is ofl«u enclosed or surrounded by 
a kind of involucrei terras] n Cupuk; such as the cup at 
W\c liiLse of the aeorn, the bur of the (.'bestnut, and the leaf- 
like cxjvering of the hazel-nut. The name Olnn* 
(sometimes Gland in EngliitiiJ is techiiicalij applieil 
to Bueb nuLs, this lieing their classical Latin namo. 
5tiH. Tlie fruit of tlie Walnuts and Ilickor}* is 
apfwrently a kind of drapaetout nut, or sutuetliing 
intermediate lietween a stone fruit aud a nut. But 
certain monstrosities give reason for BU[i|)osiu){ that, 
the seeming exocaq) (^-11), which in lliekory 
"■ hardens and at maturity debis<!es in four valves, 

is of the nature of an adnate iuvolucre. The eocoauut U a sort 
of llliro-tlruijaceous nut. 

hfi'i. Natlat^ or in Latin form Nucri^ (^Nucida), is sometimes 
superfluously employed in a literal sense, as a diminutive nut.' 
Of late it has aeqiiiretl a good and fairly legitimate use as the 
uuuie of Uie seed-like, or rather akeue-like, closed parU or lobes, 
of cnistaceuus or otiier hard testnre, into which certain bilocular 
or plurilocuhir i>criLari)« sei>arate at maturity, i. ». for the seg- 
ments of a Bchizocarp, 571. which resemble akcnes.' These arc 
Boitietiines cHr[>elH. sometimes half-caqwls, as in Vcrlx'na, also 
in itorroginaccie and Laliiutie (in which the scgmcuta are greatly 
separated in tlie ovary), and sometimes, as in Nolana. they aro 
portions of com | founded carpels which have been exceedingly 
multiplied by chorisis. 

ftT'l. There nre complete transitions between dry nutlets, with 
a thin and herlmceous epiearj), and llie pyrmtt (574) or stony 
Inner [wrlion of sueli cnqvela when dnipnoeons or composing a 
dni|ie of two or more stones. It is tlierefore a hanlly incongru- 
ous and very convenient use which extends the terra nutlet to 
include these small sire«l-likc stones also, as. for example, to 
tliostt of Holly, Hearberrj'. Hawthorn, and the like. 
071. The pair of achonium-likc or often aamai-a-tikc carpels. 

> Kui ai 

I'. lirlwKn which thprr \t no flx^l (ti>llni-ilriii, will corer 
tliit pnun'l- Tlip (ruit of Cypcrsccw, for lixuniv. i« truly %n sclieninm, 
tf thli namt' i* c^ot In be wk-A (nnd il now cnnuunnl/ ia| for any other Ihsn 
a ni«ncH-iirp<-l!ary fruit. It 1) nfien lennn) it nut, lomolimra n outlel, and 
by a )*tu wrltur, Bnx-kler. a ratTupaii. 

* Cixi'i |*inB, CnrvM. from a Gnvk word for komcl) l» anollipr name Int 
fnilt<arpi-l>. or •n«rallnic l"l»io( a dry pericarp, n» wpI I foniciiiiiceiil onea 
(of Ktipln)rl.U) ■• fiir In'lrhlKTDt. nonw •ucli Inlird nr paniMp (raili 
•re uld la bo dieetrum, Irinceoat, ic., acconllnB to the numbrr of iobci i>r 

FlO. 108. Awm mm) of www Oak. wlUi 1U rnp, or ei.p.iU 

rre KINDS. 297 

nnited by their inner face Imt sep&ratiDg entire at maturity, 
which constitute the fruit of Umbelliferte, takes the name of 
Cremocarp (Lat. Cremocarpium) ; and tlie halves are called 
Mericarfs. These names it may sometimes be convenient to 
use ; yet it is not advisable to have special names fur the fruits 
of particular families; anil mericarp is here synonymous with 
caqxd. For drj" 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- 
scccled portions, an appropriate recent name is that of Scuizocarf. 
The com^Kjncnt carpels of such a fruit were long ago named Oar- 
eeruUs {carceruli, little prisons) by Mirbcl. 

572. Fleslij Frnlts, which from their texture are naturally 
indehiscent, may be either fleshy throughout, or willi a Unn rind 
or shell, or fleshy externally and hard or stony internally. Of 
the latter, tlie type is 

573. The Ihmpe or Stone Frnit proper (Fig. 639), that of tbe 
clierrj, plum, and peach. True drupes ai-e of a single carpel, 
one-celled and one-seetled 
(or at most two-seetletl), in 

the ripening of which the / Y/sS^ A /;'■'■■ 
outer portion of the pericarp [ ^ f \i I f /. 

bec-omes fleshj' or pulpy, and 
the inner stony or crustaee- 
ous, 1. e. divides into larco- 
carp and puiamen. (o^l.) 
But the name is extended to ** •" 

pericarps of similar texture resulting fVom 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 Comus, Rhamnus, Ac. 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 dnipcs 
or drupaceous fruits of more than one cell, ripening into separate 
or separable hanl endocarps or stones, each fllle<l by a seed.' 
Bcariierriea (Arctostaphylos) and Huckleberries (Gaylussaeia) 
arc goo<l illiistrationB of this. The seed-like endocarps of this 

' Tlic term Arinus, the orifpnal name of auch a htny ns a grape, hM been 
used in descriptive bolanj for a imsll drupe or ilrupetet, and the ripened 
earpoUof Itubui hare been termed neini or ae/nM, but without discriminating 
them from berriei. 

FIO. S39. Verikul •erilnn nf a pearh. 640. An utmoncl; In wliicb tbeeiocvp, tb« 

jiortlnii of the peiinarp <li*t re|ir«Hnii< the pulp nf the pntch, i ilin Juli ilit. ud at 

length Mparatea bf datilaccnc* from tho eoiloairp, or ibeU 



sort are Ftbkh* ; and Ihe TruiU are dipyrenout, iri/ym 
IHrapijrenoHi, &c., according as they usntain two, three, or four 
pyreiiie. When the sarc'0<^ar|i is iliiii ami drit-B up at maturity, 
tliL-se |iyn-uie [>iuis by gradations into mieiiliv (itir'J) or nutlrts : 
hniee pyrenit arc not uucommonly iu KugliBli deBciiiiliuna called 
niUktt or ducu/h. 

57J. The Pome (Fig. C4l. 642) is the name oftlie apple, pear, 
and (iiiince. These are tlesLy Tniits, coinpostHl of two to 
Hcvcral caqH-ls (rarely by alxirtion only one), of 
parclimcnt-like or (in Hawthorns) bony textnr.-, 
enclosed in flesh which morphologically belong;! 
to odiiatc calyx and receptacle; as may l>e ap- 
prehended hy comparing a rose-hip (Fig. 407. iu 
fluwei-) with an appli; or a pL-ar. Of thu qiiince. Ihe 
whole flesh is calyx or h<>~|)Bnthiiim ^llii) ; in the 
appb and [jear, the inner or core-portion of the ( 
fleuh is of the nature of disk, investing the carpels. 
In the IViiit of Hawthorns, the carpels Ijecorae bony 
pyrenie (374). and so the fruit is drupaeeoiu, is 
iu<lei><l nothing more than a Kynear|KiU3 drupe. 
In Kriot>otrya, or Cnmquat, the cur|)els heeoTuing 
very thin and niemliranaccous, the pomaceous 
fViiit is in fact a kind of L>crry. 

fi76. The Pepo, or Goard-ft-alt (Fig. 64S), of which th« gourd 
and squash arc tlie tyi>e, and the melon and cucundwr equally 
familiar illustrations, is ibc char- 
aclcristid fniit of Cncurbitacete. 
fleshy internally and with a hard 
or Rrm rind, all or port of which 
is referable to the adnatc calyx 
completely incnr])oi'Blo with tho 
ovary. This is eillier one-oelled 
with three bi-oad and revohitn 
puriciai ploccnlip. or these pla- 
centte. borne on thin dissepiments, 
meet iu the axis, enlarge, and 
spread, unite willi their fellowa 
on each side, and are n'Orcted to 
the walls of the pericarp, next which they l)eflr their ovules. Aa 
the fhiit enlarges, tile BCC<t-lieBring placenta' usually cohere 
with the walla, and the partitions arc olilitt.' rated, giving tho 

ir »|iplo In irnii 
i( t,. -Itak In tlia 

la Uoonl. 841. Ungnio of on* uf 

.: tha 

1T8 Ki:sDS. 299 

appearance of a peculiar abnormal placenta tion, which the study 
of the ovary rea<Iily explains. In the watermelon the clible 
pulp all belongs to the greati}' developed placentte. Fruits of 
this family in which tlic i-ind also is soft at maturity are true 

577. The Hegperidlnm (orange, lemon, and lime) is the fleshy 
fruit of a IVcc many -celled ovarj' with a leathery rind, and is a 
mere variety of the berry. The name is 
appUcil only to fruits of the Orange tribe. 

578. The Berry (Lat. Baeea) comprises 
all simple fruits in which the pericarp is 
fleshy throughout. The grape, goosebeny, 
currant, cranberrj' (Fig. 645), banana, and 
tomato are familiar examples. The Brst 
and last consist of an ovarj' fVee ftvm the 
calyx ; in the others, eolyx and ovary are 
combined by adnation. 

579. Aggregate Fruits arc those in which 
a cluster of carpels, all belonging to one 
flower, are crowded on the receptacle into ' 
one mass, as in the raspberrj' and black- 
beny token as a whole. (Fig. 646.) They C 
may be aggregates of any kind of simple 
fruits. But when dry and not coherent, the mass would simply 
and properly be described as a head or spike of carpels, more 
commonly of akencs, as in Ranunculus, Ane- 
mone, &c. Yet when numerous carpels thus 
compacted become fleshy, and sometimes more 
or less coherent, the aggregate may need to be 
taken into account. The best name for it is 
that of SiTJCAKTirM, or in English 
form Syncabp. But the term has 
l>een ap]ilied to multiple fruits as 
well.' In Hydrastis, the numerous 
carpels imbricated on tlie upper 
part of the torus are baccate, that 
is, become berries ; in a raspberrj', 
the seennngly baccate grains are drupactoiu (being druprku, 573), 

' Thp iipicarp which i» a gyntcclum might be dealftnated a timjJr tyn- 
nir;»HM ; that which it an Inflorescence, a comjJtz tyncarpiam, which may be 
biDiir >u«. pautifloroUB, or multiftoroua. 
FIO. m.t Tlie iBtger Cnmbetrr, Vacdnlon 



and, slightly cohering leather (though without oi^nic nnim), | 
they lill as one body from the conical dr}- torus at maturilj-. Jt 
is the same in blackberries or hranible- berries (Fig. C4C, G47), 
except timl the drupeleta [jorsist on the torus, which jjartakes of 
the juicincBs.' In the agga'gate fruit of Magnoha (Fig. 648-650), 
BUcU car[wls, imbricated over one anollicr, cohere more or lesB 
at nil contigiiouB iMivts, aud 
become drupaceous ; nci-cr-. 
thcless, at maturity each 
opens doreally, allowing tlie 
seeds to fall out : in age it 
dries and hnitlens, and also 
f>eparat«s from its connec- 
tions, and so be- 
\ ^ comes a folliclo,but 
ivith the remark- 
able peculiarity gf 
\ dorsal iustcad of 
ventral debisivncc. 
/ (Fig. lioO.) In Li- 
riodrndi'on, s tree 
of the same family, 
h eariH^ls arc 

dr}- and indehiscent throughout ; and they lai^'ly consist of long 
and flat styles, iiubriialud in a cone, but separating from each 
other and from the slender tonis at maturiti-, when each becomes 
a sumara. 

580, AcceaaoTj or .Inthocarpons Fniits arc those of nliieb some 
conspicuous portion of tlic fhictiflcatiou neitlier belongs to the 
jlUtil 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 Pieudociirp, This 
condition may occur either in simple, in aggregate, or in multiple 

1 Tho Bpcrrjinte fruit like llml of Rubua (named by tonw VouacnrpiHm, 
by othenon .Etrria. Ergtkrvtlomum, Sc.) wa* l(.'nn(.il by l)unii)niiT R I)nipt- 
larn. A (Imilar nggregnlion of luiL-cate carpels he l[mir<l a /junrfan .■ iit 
folliclca, a FaBiirhm, &c. All such namps may l(K>k well iii a «yeli'lil ; but 
llicy are both (uperflnoiu and iinmanagcnbU' in phytograpby. 

PIO. DHL AEgTKiaU rrutt of ['mill 
whI fhm a tnmrt <1pl>lM«nit nriwl liiiii 

<tAtiiiilKil, nt f>ill ButlntUjr, <lrl«il up, ilomllr ikli W i n it. upadng Ilie pak of Moti of 



581. Gaultheria procumbens, the aromatic Wintergreen (Fig. 
G51, 652), affords a good example of the first. Its seeming 
beiT}' (the cheekerberry) , with summit crowned by the tips of 
the calyx-lobes, well imitates the true 

berry of a Vaccinium, such as that of - "jnciMD: 

Fig. 64 5. But it comes from a flower /2^\ I ^S ^K 
with thin cal^x, underneath and free iJiKvii^ 
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 fniit, yet 
without adhering to it, and in ri[)ening counterfeits a red 
berr}'. So in Shephenlia, or Buffalo Berr}', the seeming sarco- 
carp of a dnipe is i-eally 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 calyx, 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 Blackberry 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 disjiersed over the surface being 
apparently insignificant. Equally in many ^ 

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 combine<l, as in a pine-apple. 

583. Multiple or Collectlre Fmits ^ are those which result from 
the aggregation of several flowers into one mass. The simplest 
of these are those of the Partridge-Berr}' (Mitchella, Fig. 465, 

1 Collective is the preferable name. Tlie term mM/Je was applied by 
DeCandoUe to what are here (following Lindley) called atjffrtfjttte f raits; 
and the cufgregate fruits of DeCandoUe are here called multiple or coflective. 
Moreover, the distinction between accessory or anthocarpous and collective 
or multiple fruits was not recogniied 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, Aooreoati ; 

FIO. 651. Forming capsnle of Oanltberla procnmbens. with enlarging calyx partly 
covering it. 682. Same, more advanced, and In longitndlnal sertion. 
FIG. 653. Vertical section of half a strawberry. Compare with Fig. 406. 



4GG) and of aome species of Honey sucIHe, formed of the o 
of two blossoms united into one flesliy fniit. The more usual 
Borta are siicli as the pi ue- apple, miiibeny, and the flg. These 
are, in fact, dense fornis of inDorescencc, nitli the fruits or floral 
cnvelojKS mattwi together or coherent with each other ; and all 
or some of the parts succulent. The grains of the mulberry 
(Fig. ()o4-iiiHi) ai'G not the ovaries of a single flower, likn those 

of the blatkUm winch it wpcrflciulh i-esembles: thev Iwlong 
to as mnny sepal att Howcra and the |)ulp i»crtains to the caljx, 
not to the jwncarp whi(.h is an akcne So that thifl. like moflt 
nuilHpl(! fniits la antliocarpous as «tll as multiple. Similarlj', 
the mostly indcRnttc fVuctiferous masses of htrawbeny Blitc may 
resemble strawlwrries ; but the pulpy pait is the call's of many 
flowers, not the suceulent receptacle of one. In the pine-a|}ple. 
tli« flowers are 8]iieftlc or capitate on a simple aiiis. which grows 
on Iteyonil them into leafi' stem ; this when rooted as a cutting 

9. Frujl Fomponntl (oTarla compounil), Stucakpi ; 4. CollMtivc fruits, 

Later, hi W« " Eleiticnti of Botuny," Linilley rwluccil the cUmw ta 
fno: I. Simplr Fmili. thosi- proceeding from a einglc Bower; 3. itnU^Jt 
X""'l'i ibose formed oul of teveral flowm. 

ria tM. A n>iillwrr 

n IiMllliUr Mnwrn vlih Be 
niKllE W>. t>n>iiil 

II. Tha* 


bears another pine-apple, and so on ; the constitoent flowers have 
tliroiigh immemorial propagation in this v&y become sterile and 
seedless, and all its paits, along with the bracts and the a^kis of 
the stem, blend in rii^ening into one fleshy and juicy mass. Few 
fiTiits of this class have ever been technically named, at least 
with names which have come into use. But the two folton'ing 
deserve si^ecial appellations, although only the latter is familiar 
either in ordinary language or in descriptive botany. 

5S4. The SfCODlam or Hrpontbodlnm, the Fig fruit. (Fig. Go7- 
G59.) This results from a multitude of flowers concealed in a 
hollow flower-stalk, if it may be so called, which becomes |>ulj>y 
and edible when ripe ; and thus the frut't 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 ver\' small akencs, which are commonly taken for seeds. 
The fig is to the mulbcrrj- what a rose-Uip is to a strawljenj*. 
(38'J, Fig. 406, 407.) It is further explaineii by a comparison 
with a near relative of the Fig-tree, Dorsteuia, in which similar 
flowers cover the upper surface of a flat peltate disk. This disk 
or plate sometimes becomes saucer-shaped by an elevation or 
incur\-ation of the maigin. 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. 

58o. The Strobile or Cone (Fig. GCO) is a scaly multiple fhiit, 
resulting from the ripening of certain sorts of catkin. The name 
is applied to the fruit of the Hop. where 
the lai^e and thin scales arc bracts; 
but it more esi>ecially belongs to the 
Pine or Fir cone, the peculiar fhiit of 
Coni/era) (507), in which naked seeds 
are liome 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 as 
that of Cypresses, 
has been termed a "" 

Galbuxus, an unnecessary name. The galbulus of Juniper is 

hubeen iletacbfld. 



remarkablG transformation into a seeming beny ; the C 
cohering with each other as thcj' gron and b«ooiuing fleshj o 
maturity, conii)lettij" enclosing a few bony-coated seeds. 

o8C, A .SynupBiB of the kinds of Fruit, as chai'act«nzed i 
this ehapttT, is appended. The analysis extends only to simpla 1 
IVuits. For there are no commonly used special names of , 
kimis of Aggregate (579), Aeceaaory (080), or Multiple (583) ' 
ft-uils, except that of Strobile. 

Dry Mid dchiii^mt, rnonocarpelliir/, 

Opcningby one (uhitfl; the ventral) •"^'"^ Fot.ucLK. i 

Opi'TllDit Lijr bulli sulurea, LEoDMit. 

Or tni[»vcrM!l}' Jolntnl, Lohimit. 

Dry and iIvhJKcnl, bl-pluri-carpcUary, Camulb. 

Wlii-n Itl dehiscence is eircumseiieile PVKl*. 

When dchiic^ni by two raltc« from two parietal pUL-cntv. . . Siu(|i;b. 

A short and hruad siliquc, Siucui. 

Dryaiid bt-plurl-CBrpcllary.iplillinginloone-H-ededcarpeU, . ScnizoCABr. 

The diiuemiu jrhiwwarp at UtobelUtL-r* CuBHOUAar. 

Eni^h at iu halvn or carpeb Hehicarp or Mehicakt. 

The akene-likc or nut-liku parts into which Schiiocarps Kcorrally 

divhle SucuLBs or Nuil«¥«. 

Dfy and lnd«'IiliictDt. one*eIled, onc-lw(«ecded, 

Winged SaxaKju ' 

Winglcw, and with Ihe 

Tliin pericarp eoniolidated with the seed CABtopsis 

Thin pericarp loose and not fllled by the seeil Ctbicle. 

Thiutt or hard pericarp free from tlie seed, 
Small, from a one^elled ane-lwiM> ruled ovary, Akeitk orAcBWiiirK. 
larger, nioslty from a iwo-scveral-celled and nvulcd ovary, . Iter. 

Nul borne in a eupule or involucre Gt.aita. 

Fleihy and indehiscenl. | 

Heterogeneous in texture, having 
A stone (puiamen) or ntillets wHhln an exterior »«reoc«rp, . DRura. 
Papery or curlilaginnus carpets in an inferior sarcocaqi, . . PDklB. 
A harder or firm rind or exterior, and soft interior, 
From an Inferior ovary (eonflned to Gourd Family), . . . Paro. 
From a superior ovary (confined- to Orsngi' Family), IIesperidicv. 
Homogmcoui, fleshy tlirougliout, Bkkht. 

THE SEED. 305 



587. The Seed is the fertilized ovule (515), with embryo 
formed within it. It consists, like the ovule, of a nucleus or 
kernel, enclosed b}' integuments. The seed-coats are those of 
the ovule, viz. two, or sometimes only one, in certain plants 
none. Occasionally an accessory coat apiKjars after fertiliza- 
tion ; and certain apiK»ndages may be pnxluced, as outgrowths 
from some part of its surface or from its base. The nucleus or 
kernel is composed either of the embrjo alone, or of a nutritive 
de|K>sit in addition. (19-41.) All the imrts of a seed are in- 
dicated in Fig. 6G3. 

588. The Seed-stalk or Podospekm, when there is one, i% 
the funiculus of the ovule (516), and retains this name. So 
also do the Ciialaza, Riiaphe, and IIilum ; the latter being the 
scar left by the separation of the seed from 
its funiculus or directly from the placenta. 
The foramen or ovule, now closed, is the 
Micropyle of the seed. 

589. The terms which denote the char- 
acter of the ovule, such as orthotropous^ 
campylotropou8^ amphitropou$s and anatrapoiu^ ^Ppl}" equally to 
the resulting seed. 

590. Seed-Coats. The integuments of the seed answer to the 
primine an<l secundinc of the ovule. The main seed-coat is the 
exterior integument of the ovule when there is more than one. 
Being the most frnn coat, and not rarel}* cnistaceous in texture, 
it takes the name of Testa, which is equivalent to seed-shell. 
It has also l)een named Spermoderm (seed-skin), and sometimes 
Epi sperm. The latter name (meaning xi]yon the seed) is best 
applic^l to the iK»llicle or outer layer, sometimes a thick one, 
which the testa of c*ertain seeds forms. The testa is extremely 
various in fonn and texture, is either close and Conformed to 

no. «13. Vi»rtl<-nl mairnificd nertlon of the (anatmpouH) neeil of tho American Lin- 
den: with the parts inilk^tod, viz. tlie liUum (a); testa {h)\ Xff^\^r\{r)\ aUniroon (</); 
emltnn {e). w\. Vertical section of Uie orthotropoas seed of HelianUiemam Cana- 
dense, witb its ftinlculas, cu 




tile nufleiiSi or loose ami cellular (as in PjTola-seeds) , or rari- 1 
ously nppendugwl. 

!i'Jl. Tbo iiiuer coat, coUwl Tegmes ami sotnettnics Ekimj- 
PLEfRA, when present is ulwaye coulbitneil to the iiuvlcus, and 
is thill or soft and delicate. Somctimea it is inconspicuous 
through cohesion with the nucleus or with the inner surface oT 
the testa. In ovules of one coal it is necessarily wanting. 

dS2. Appendages or outgi-owths of tlic testa generally Iiai-e ' 
reference to dissemination. Two chavactei-istic kinds of such ' 
appendages arc the wing and the coma, ' 

both i>ertaiuing only to tlic seeds of dehis- 
cent fruits and calculated, by renderiii;; 
seeds buoyant, to facilitate dispersion In 
thcwind. The wing of a Pine-scwl (V\u. 
CCl, 602) is a part of the cariwllarj- sinli 
upon wliieb tlie two ovnles grew. In 
Trumpet Creeiwr (Fig. 605), an enlui' 
wing surrounds the body of the snil 
In the related Catalpa (Fig. G6(i), ii i- 
niniol}' extended ri-om the two ends, iiii'l 
almost dissolved into a coma, the tian 
given to the tuft of soft hairs like i!. 
which foims ILe down at one end ol' i ' 
seed of Milkweed (Fig. GG7), anil ui 
K|iiloliium, and at Ijoth ends in several 
Aijocynnccie. In the Cotton-plant, very 
long and Sf.ft hairs, admirably adapted for 
spinning, Lliiekly cover the whole sccd- 
■" coat. The wing anil coma of seeds are 
fiinctionally identical with the wing and llie pappnaof the pericarp | 
in the samara and the akenes of Compositte (.'iG3, AG'l), but I 
morphologically quite unlike them. 

5113. There are other (mainly microscopic) stnicturcs on a 
seeil-coats which come nseftilly into play in arresting farther ' 
dispersion at n propitious time or place. In many lint not all 
Polemoniaocte (notahly in Collomin), in certain Aivnthacvn, 
eoch as KueJlia tulierosa (and equally in certain Conijiosilie of 
tlic Senecio trilN> nnti in Sahias, Ac, among Labiatx*, where j 
tliis stnietnre is transrcrrcd to akcncs and iintlets), the testa is I 
coated with short hairs, which when wetted burst or otherwise 1 
njien and discharge along with mucilago one or more verj' att«a- I 

PIC. MO, Wlnp>lHint»fTruni|MtCrMpiir.Taonin*r«lk'B>i 

a TllUofCatlllMi, i 



iiuted long threads (tpirielet) which were coiled within. These, 

protruding in all direttiona and in immense numbers, form a 

iimbus of considerable size around the setd, and evidently must 

serve a useful end in fixing these small 

and light seeds to the soil in time of 

rain, or to moist ground favorable to 

germination to which thej may be 

carried bj the wind The mucilage so , 

laigely develo|je<l iiiwn the seeds of 

Klax, Cress 4.c when wetted 

is probabh of similar use 

394. nhile the testa in 
many seeds is hard and 
cnistaceous or bonj imitat- _ 
ing the pcmarp of a nut in othora (such as Fteonia) it becomes 
berry-like (baccate) and in Magnolia drupaceous ^ (Fig 0( 8- 
671.) These ma\ also be r^arded as adaptations for dissemi 

nation, here bj the agencj of birds, attracted bj bnght colonng 
and edilitc pulp. 

:)95. The riiaplic of an anatropons seed (shown in Fig. C81. 
685) is sometimes so salient as to form a conspicuous n|)|>on- 
dage, as in Sarracenia, Fig. C72. Again it may be whully 

■ Si-c arlivle On the Stnicturc of the Ovule and Sood-conts nf Magnolia, 
in Jour Linn, Soc. ii. 100, from which the accompanying flguret and Fi^;. 
58!t-0Q7 are rcpnuluced. 

FIG. 6W, FurmluK smxI (niM djflith nf an Inch tfngt nf Mapinlln rmbrelt*; Mie 
rtiiLi'he (nw!irrl tliR ey. em. yiagiMvt rltw nf tho nmr clltl'lv.l lenRtliHlK I1irniii:)i 

■ILMIii.'I n»in ItilM b> Ihcinnrr tnttit), Iniin«llat>lT«i<;l<«lTig tlia nuclenR. r. Tlienpjm- 
•ilf iO-Ib i-t tlw loU ti tliU'lur nn account nf the rluiphe, In wlikh d liKltcatH die cwl 
of rjinl •IncI*. 

PIO. an. Anear1]r rilll'icmnKciLnftherwtnraliriM «7I. Lmidliirthuil FMIInn, 
MilnrKwI. rtmirliig thecniiiiwwnunr >tonr InncrMratiim nf the tnrta mil ilevelnped : 
the t«iti IgltenU at In Fli. MO. Gil. A transTenaicctlan In IbeMDMiodUon. 


inconspicuous, as in the ri|)o seed of Magnolia, wlicrc It*^ 
length coni|ileleIj mfrged nncl imbeUdeil in tlit licsliy (lru|>a 
testa, as shown in Fig. 670-G72. 

oUG. Crcst-Uke or other appcudages are not uncommon uitder 
on llic rhnpbe or at the hilum. These are outgi-unttis )tnMliicecl ■ 
during the development of the ovule into the seed. Id Ssnguf- I 
□aria, Bucli a crest develops fVom tho whole leogUi of the rUai»he 1 

(Fig. 073) . in Duentia, Coi'^dahs (Fig. GTi), &c.. from soma j 
pait of it, mostly from its base next the hiliim. or ftom tha ] 
liihnn itself, or even fVom just below it. Such an nppcurlnge, 
es{)ceinlly when ottaehed to the base of the seed, is naniPd a I 
Stkoi'uiole, a similar and commonly a wart-shaijcd a[)pentlage I 
in EupliorI>ia, Iticinns (Fig. 675), &c., is produced by an out- I 
growlli of the external orifice of the ovule, the micropyle of tho I 
seed. This properly takes tlie name of Cari'nclk. Bnt the I 
two terms are not alwa^'s discrimiunted. By further develop- I 
mcnt, cither of these may give rise, in certain seeds, to an acce»- I 
Bor>- covering called 

397. The Aril or Arlllas. This tcnn, rather vapnely employed I 
by Limucus, was flret well defined by Gtertuer. The true ariUttt J 
is an accessorj' seed-covering, more or less incom- 
plete, formed between the time of fertilization and I 
I -. « the riiK'ning of the seed, by a growth Ttoro the aper J 
VriT of the fiminiUis (when there is any) at or Just be- I 
U^/ tow the liilum, in a manner similar to that in which J 
V"i tlie coat or coals of Ihe ovule are formed. That I 
"" of Nymphiua (Fig. fi76) is a typical example ; only J 

the arilhis is devclojicd n'om the Ainicnlus at a ]>oint distiocdy f 
below its apex : here a ring forms, which grows into a cnp, and 1 
tills is soon extended into a sac, loosely enclosing the seed, i 
Open at the top. This is membranaceous : commonly it is fleriiy. | 
When there is aiwoliitcly no fiiniculus. the aril may originato 1 
from the [>la[.-enla, as it does in PodophylUini, in which most of f 

FIO. ItT! An«ln>tMuiiiiBeilnf SsmuamU pufpuren. wllh very «1l<iil ih«I>l». m. I 
8iUBpnr8nngil1n»rl»nt Blnoilnnl.wllh rliaphrpnwlwl f'T It! wlmli. iM.fili (tl*. 
of Curyilalb aunn. wttb cmt ur >lmphlula. ulUfbad »l or luv Uie lilliim. STS. 
(■mpamlcd) sf Itlrlniia, wtcli t>> nmnf 1(. 

fiti. STd 8«c>lurW1ilicWiiict'LII)'.Kyuiiil>Eaoiluntla,lii ItaluiKUiiilliluU 


the pulp of the berrj- consists of these fleshy arils, much com- 
pat-ted. (Fig. G77, C78.) 

598, The kciniate nril of the nutmeg (mace) and, it is snid, 
the biiglit red and pulpy aril of Enonymus and Celostrus begin 
in the manner of a ca- 
runc'lo, and are formed 
(mainly if not wholly) 
of nn outgrowth at or 
around the niicropyle. So 
that, if an orthotropous 
seed ever develoi>ed an 
aril of this sort, it would 
be seen to begin at the aj^x of tlie seed and cover it fVoin above 
downward. I'luiK-hon, who distiiigui sited this fVom the tnic aril, 
gave to it the name of Akillode (Arillodiam) or False Arilhis. 

50y. Tbe Koclens, or kernel of the seed, consists of the Albu- 
men, when this substance is present, and the Embryo. 

COO. The Albumen, as described in the second chapter (25, &c.) , 
is the name generally employed hy systeniatic botanists for a 
store of nutritive matter in the seetl outside of the emhrjo, 
whatever its chemical comi>osition. It is not here the name 
of a chemical substance {nlbumen or albamin), but of a cellular 
structure, tire cells of wliitb are loaded commonly with stuivh- 
graius (as in the Cerealia), more or lees mingled with other 
matters, or else filled witban encnisling dejxtsit of some equiva- 
lent substance, as in the cocoanut, coflce-grain, &c. Tlie cells 
in which this deposit is maile belong either to the original tissue 
of the nucleus, or to a new formation within the embryo-sac, 
mostly to the latter. (503.) 

601. Albumen may be said to belong to all seeds in the grow- 
ing sta^rc. In what are called albuminout seeds it i}crsists and 
forma cither almost the whole Itcrncl, the embno remaining 
minnle (as in Fig. 23. 54, 680), or fonns a lai^ iwrtion of it (Fig. 
l;t. 17. 10, 21. 48, 663. CG4), 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 disapjiear 
altogether, as in the seeds of Maple, Almond, Squash, I'ea. and 
the like, which arc therefore said to l>e exalbuminoiis. The 
difference Iwtween albuminous and exalhuminous seeds is that 
the maternal nutritive dejxisit is transferred to the embryo in 

FlCi. TTT. Sn'tlim nt perlMup nnil p-Incenlii nf Pnloi 
Med detactitii and eDbtrg«ii, iUtIiIhI lenSiliirUe, (bowttij 

310 THE SEED, 

the former during gennitiation, iu the tatt«r during the g 
of tliH sL-ed. 

C112. The albumen was named Perispeim hy Juseieu. and I 
Eadutperm by Richard (2a, note) ; but neither name has in | 
sjstenialit! botany clisplacocl the earlier one of Grew and U:ert- 
iicr. Uiit both names liave I'eeently been bruuglit into use tu 
distinguish between two kimU of albumen, that formed within 
tlic cmbrvo-sac. whidi is siieciSeally termed ENDO^rcoM, and that i 
foimnl withaut which take* the name of I'erispekm. This use 
comports wilh the etymology of the two wonls, the former refer- ; 
nng to a comparatively int«mal and the latter to an external 
portion of the siwl or kernel. 

603 In most seeds the albumen is endos]ierm: in Canna it ' 
IS nil peniperm In Xymphiea and its allies (except Nelum- 

1 iiira, whifh iia8noiie)mostof itisperiBperm; 
but a thin and condensed layer of endosperm 
I surrounds the embryo, where with the pcr- 
isitent embryo-sac (or the ajMsx of it) it j 
t irms the fleshy sac in which the embryo is 
I nelosed. It is the same ui the Pepper Family 
(Fig. 679), except that there is a latter quan- ' 
tity of endosperm or inner albumen. 

604 \\ hen the nucleus of a ripe seed is hollow, as In tlie 
cocoamit and nus Nomiea, tlio formatiou of endusponn, which 
usualh begins next the wall of tlie embryo-sac, has not proceeded 
BO as to nil the cat itv. The embryo-sac in the cocoanut attuus 
enormous size, and the cavity is filled by the milky fiuid. 

fil}5. The lextni'e or consistence of the albumen differs gr9atlj. 

It is Jarinaceout or rofo/y when, consisting mainlj- of starcli- 

groins, itmay readily be broken down into a powtier, 

as in wheat, buckwheat, &i: ; oil*/, when saturated 

with a fixed oil, as in i)oppy-9eed : Jlethy, when 

more oompacl. but readily cut with a knife, as in 

the seed of Barberri' ; mueilaginou», when soft and 

somewhat pulpy, as in Mnrning Glory and Mallnw, 

but when rlrj' it becomes Seshy or harder ; corneous, 

"' when of the (e\ture of horn, as in cotfe* and the j 

Boed of Canlophylltim ; and e\en boni/. as in the vegeUble ivoi^", 

the seed of Phj-telephas. It is mostly uniform : but in the nutro^, 

no (Wi. t.<inKl"r> 

ml ot tbfl Ki-oLlai] ritptw, J 



in the seeds of Asimina (Fig. 680) and alt the Custard- Apple 
family, it is marked by traosverse lines or divbions (eausetl bj' 
iuflexioDS or growths of the inner seed-coat), giving a section 
of it either a marbled appearance, or as if it had been elit by 
incisions : it is then said to be ruminated. 

COG. The Embrjo.' being an initial phintlet or individual of a 
new generation, is of course the most imimrtant part of the seed. 
To its production, protection, and supiH>rt, all the other parts of 
the fruit and flower are subservient. 

COT, In an embryo of full development, namely, one in which 
all the parts are manifest antecedent to germination, these parts 
are the CaulicU, otherwise called RadicU, the Coli/ltdimt, and 
the Plumule. (20, 30.) The first is the initial 
axis or stem, a primary intemode ; the second 
consists of the leaves of tbc primarj' node ; the 
third is a bi^inning of a farther growth which 
is to develop more stem and leaves. Such an 
cml)n-o is usually imaccompanied by albumen, 
having in the course of its growth taken into 
itself (mostly into the cotyledons) the provision « «» 

which in other sec<l9 is mainly accumulated external to it until it 
is drawn uj^n in germination. 

1 The word Fmhiyo or EiBhryrm wii« spplied to Ihii hoAy In plums by 
Bonnet {ConsidiJrK lions Bur 1m Cnrpi ortwniB^cs), In 17«2, and w>a inlmdn(-o<1 
into Bjilomaliu Ixitany iit aliout the snme time (IT63) by Ailnnenn: It wm 
taken up by Gsrtner in 1788. Juuieu in the Genera Plantarum (l~^) li^ld 
to the term CornUm (the cor tenii'mM) wliich earns down from CBsalpinlw. 

Brinfc llie ^rniinal part of Ibe seed, the embryo of the plant, like that of 
the animal, is in general language often called the Germ. 

FIG.esi. Sml nr ■ violet Un>itm|ini»X snlarmKl: witli hMani or mr { 
b). ukI dialiua (r) Indtntral. e»s. Verllnl Bctlon of tlie umc, iliowlng tli 

FlU. (W3. VeRI<i1 iwtlnn nf Ili« (nrllialropoiu) reed ol Buckwhsat, iL 
miliryo fnlilsl mnml In the mnlr albnmcn. 
Fin. DM, Vartlral ivrttnn of the (aiiMnitxHU)Md of Elodea Virginia, tl 


nnletflv BlUnR Hie 

plilnlnm triwrne (anaimpomi 
Iteral ■• In F\g mi. Cm. Vert 
, e, the legmen,/, tbe albanien 

312 THE SEED. 

60S. The opposite extreme is an embryo (as in Fig. G6C) 
wbic-h appears as a moi-e speck in Ibe allmmen. but iti whiob 
ciosc microseopipal ius|>ect]un may eonimoiily reical wjrni.' differ- 
entiation, sucli as a sIio:l]t noti-h at one enil (that farthest re- 
moved fromthe micropyle) ofa tlicotjledoDona embryo, indieatin); 
the future cotyledons. Indeed, in Monotropes, Orobnnchat-eje, 
and some other parasitic dicotvledonona plants, and in Orchkls 
among the monocolyledonous, the embr^'o is a globular or oblong 
partii^le, with no odumbratiou oT organs whatever antennlent to 
gennination. Tliere are all grades between the most rudimeti- 
tary and the most developed embryos. 

6119, Cnder tlie circumstances of its formation (aS2). the 
radicular end of the euhr^'o is always near to and points towards 
the micropyle of the seed, viz. to what was the orifice of tie 
ovule ; and if the embryo be straight, or merely i>artakes of the 
curvature of the seed, the eot,vledons point to the opposite 
extremity, tliat is, to the chalaza. 

GIO. The position of the radide as respects the hilnm raries 
with llie different kinds of seed. In the orthotro|K(us form, as in 
fleliantlienniin (Fig. 664) and Pepper (Fig. 679), the radicle 
necessarily points directly away from the hilum.' In tlie anatro- 
jjons foiTO, as in Fig. 663. 682, and 684-686, the t-xtremity 
of the radicle is brought lo the immediate vicinity of the hilum ; 
and so it is, although in a different way, 
in the campy lotropona seed (Fig. 689, 
690) ; wlille in the ampliitroi>otis the 
radicle |>oiiits away from the liilum latcr- 
« "» ally. As the nature of the ovule and seed 

may nsually lie ascertained by external inspection, so the situation 

' Two IccIiniuKi terms, early inlrodueeil by Richaril to iiiilicBtc (lie direo- 
lion of thf rndlolc ((■uliile). or railier its r^laiinn lo tlie liilum, «rc 

AfiiiTopoa; whwi the embryo din«ts iu radicle away from tile lilluni, u 
it mnat in all orthulmpoiu let^di; 

OrbWro/MHi. alM) iumatropoia. when directed lo the hilnm (mnre ilrlclly in 
tbe micropyle close tu the hilum), ■> in analropou* loedi. These two ivmu 
■K ilill einpluy<!d by many botanitl*. although tuperfliioiu when llii- ovule 
or iwd ii itatcil to be antln>poa« or orlholmpous, &•;. And the ttrm 
ortholropauB. ao vkA, is liable to be confiucd with ortliolropuiu as appUvd 
to the DTuIe. 

Kit-hiirtl, mnrmvcr, Icrtneil the enihryo ow/iAiVrojioiu whpn curved or mhIkI, 
on in Chiiikwi^ (Pi|[. (t8fl| and nil «ach nmipyloiropou* nci'cls; and Artnv. 
IroiiiHu when neithi? rulk'lii nor cotyledons point to the hilum, *> ocvura 
ill the icml-Bnairnpna* or amphitropout ovniv, Many butaniett ileicril e 
liic hut by the csprrulDn " nuljuir vague," or, belter, '■ embryo iranivcrae." 

Fia. aw. riun|>>1<>lr'>|i'KiB tnvl nf nnrnnim Clilc1iw»-I. magnlBed' aM Soctlon of 


of the embr^'o within, and of its parts, may often be inferred 
without dissection. But the dissection of seeds is not generally- 

G 1 1 . The direction of tlie radicle with respect to the pericarp 
is also noticed by systematic writers ; who emploj' the tenus 
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 vague^ 
when it bears no evident or uniform relation of the kind to the 

012. The position of the embryo as i'esi)ect8 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 Gmins (Fig. 5G-G1), in Polygonum, 
&c. When external or nearly so, and curved, circularly around 
tlie albumen, as in Chickweed (Fig. 690) and Mirabilis (Fig. 
17), it is said to be peripheric, 

613. The embryo maj* 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- 

Gil flBS flB8 eM eas 

volute from the a^^ex, as in Bunias ; or else doubled up and thus 
hiplicately convolute^ as in Sugar Maple, Fig. 2. Two modi- 
fications ai*e more common, and are of such classificatorj' imix)r- 
lance in Crucifene as to need special reference. Namely, when 
cotvledons 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 
cotyledons are longitudinally applied to the radicle. These 
ditferences were first employe<l in the classification Cruciferai b}- 

FICi. 691. Convolute embryo of Calycanthnii, the upper half cut away. 

FIG. 692. Seed of a Cruciferous plant (SIfiymbrium), with incumbent cotyIe<lon8, 
dlvi<lc4i. 693. Embryo of the fv»me detached entire. 

FK;. C94. Seed of a Cruciferous plant (Barbarea) with accumbent cotyledons. 
61'5. Tlie embryo entire. 

314 THE SEED. 

Robert Brown, and were adopted as priinary and tiibal cbanoten 1 
bj- I>et'andollc. 

G14. As to number of cotjledons, the two types of embrjo 
are the 

Munocotyledonnua, with a single cotyledon, i, t, leaves at the 
first nodes alternate (3!l) ; and 

DieolytedtmouB, with a pair of cotyledons, i. e. leaves of the 
first node in the most simple whorl, a pair, in other words, op[K>- 
site (il) ; with its roodifieation of 

Poiyculyltdoiioui (38), the leaves of the first node in whorls of 
three, four, or more. This occurs with eonstancy in a majority 
orConirerK(Fig. 48,49). oecasionatly and abnormally in sundiy 
ordinary dicotyledonous siwcies. 

(ilo. There are several embryos of the oolyledonous tj-pe in 
which one cotyledon is smaller tlian the other, viz. the inner 
one when the embryo is coiled or folded. And in all the specie* 
of Abronia (a genus allied to Mirabitis, Fig. 18) this cotyledon 
is wanting, so that the embryo becomes techuieolly tnonoootyle- 
donous. In another genus, the Dodder (Fig. 78, ~'i), both 
cotyledons arc constantly wanting ; and the plumule shows only 
minute scales, the taomologues of succeeding leaves reduced 
almost to nothing. 

616. Sometimes the two cotyledons are consolidated into ono 
body by the coalescence of their contiguous faces; when thej" 
are said to be eonferraminale. This occurs more or less in the 
Ilorsechestnut and Buckeye (Fig. 41. 42), and is striking ia 
Oie seed of the Live Oak, tiuereus >-irens. 

617. The general mori>hology of the embrjo and il« develop- 
ment in germination were described at the commencement of tiiis 
volume. And BO the eompleliun of this account of plant, flower, 
IVuit, seed, and emhrjo brings the history round to the starUng 
{x)int. (IS-t'J. &(;. ) Having mastere<l the morpholi^y and 
general structure of the higher grade of plants, the puiitl may 
go on to the iiior|ihoI(^y and structure of cells (or V'egetabUi 
Anatomy or llisti>l'igy). and to the study of Cryptogamooa 
Plants in all their grades. 




Section I. The Prikciples of Classification in Natural 


C18. Taxonomy, fi*om two Greek words which signify ari'ange- 
ment and law, is the study of classification. This is of utmost 
importance in Natural Historj', 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 coiTcct, will be an epitome of our knowledge of plants. 
Arrangement accoixling to kinds, and of special kinds under 
the more general, is common to all subjects of stud}'. But the 
classification in Biological Natural Histor}-, that is in Botany and 
Zoolog}', 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 he questioned whether or not they came from common 
parents or near anc*estors. 

C2(). 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. IndiTidoals are the units of the series which constitute 
species. The idea of individualit}' 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 comp>onent 
parts are reciprocally means and ends. Individuality is a main 



tUsliiic'tion between beings and tilings ; but, nlthough the t 
ency to indivijuatioii begins willi life itself, it is coaii>leti;Iy ! 
realized only in the liiglier animalB. 

6*22. la plants, as also in sumi; of the lower animuls, iiulividtt- 
alily 13 merged in cominmiity. No plunt (ext-ept one feduccd I 
to tlio siinijlicity of a single cell, of cii-ciiniscrilK^l growth, aud 
wiUiotit ui'gaiis) is an individual iu the Beiise tliat a mun or a I 
dog is. (16, liiC.) Tho lieib, shrub, ajid tree are neither J 
hidivisiljle nor of deflDite limitation. Whether their sutx^saive | 
gi-owths are to remain parts of the pre\ion9 plant, or to he iiide- 
pendent plants, deiJCixls u|>oii eiix;umslauees ; and there is no 
known limit to budding pro|>agation. 

G28. There is, however, a kind of social or coriwrate indt- 
viduaJity in those animals, or eommunities (whieliever we eatl 
them) of the lower grade wliieU are multiplied by buds or orF- ' 
shoots as well as by ova. anil in which the offspring remains, or I 
may i-emahi, oi'ganieally eonnetl<?d with tlic stock. The poly- 1 
pidoiu or iwlypariimi eommonly lias o certain limitation and a j 
dclinilc foim ; and ceilain polyps may liecome organs witli | 
apecinl ftinetious subordinate to the common weal. This is 
3 largely tine in the vegetable kingdom. So that for de- 
scriptive purposes, and in a just although somewhat loose sense, i 
the herb, shrub, or tree is taken as an indit-idual. But only 
while it forms one eonneeled body. Offfehoota when separately 
estahlishcd are eqnally individuals in this sense. 

C24. What it is in plants whitih philusopliic^Uy answers to tlie 
individual in tike higher animals is another queation, to whEcli , 
s ansnei's have been given.' Some insist that the wliole ^ 
vegetative produtt of one seed makes one indi\i(hial, whetUei' 
connected or separati^l (as may happen) into a million of plants, 
but a common and less strained view rcstriels the Inilividunl 
to sneh pi-odnet only while organically united. Othcra (of 
which Thouars at the beginning and Braun at the middle of the J 
present eentni'y ai* leading examples) take each axis or shoot | 
with its foliage to i-epresent the indlndiial, of which the leavi 
and theh- homulognes are organs, the branches lieing usually j 
implanted njKjn the jtaivnt axis as this is implanted in tJie soil, 
bnt also eqnally capable of jjrodncing roots by which tliey may ] 
make their own eonncL'tion with the soil. Still othere, on pr©- J 

> For Ilic hittor; of n[iintun upon niid a full prc^i-nlation of llil* I 
KC Ak-iLtuiiln- Bratin'i Meniuir lurieiiuillv publuliiil in ilir' Ablian<ll, Akad. j 
Wi«n-n«cliafitn mi BiTliti. IMS). ])a> In.llvlduum iUt Pflanri-. *e., a 
trmislBtioti liy C. i'. Hlone in Amcr. Jour. Se\. act. 2. xix. xx. \>^. 


cisely similar grounds, carrj' the analysis a step farther, and 
regaixl each pliytomer (IG) as the individual. Finallj', some, in 
view of their potentially indei^endent life, take the cells, or units 
of anatomical structure, to be the true individuals ; and this 
with sufficient reason as regards the simplest eiyptogamous 
plants. Uix)n the view here adopted, that plants do not nsc 
high enough in the scale of being to reach true individuality, 
the question is not whether it is the cell, thephytomer, the shoot, 
the tree, or the whole vegetative product of a seed which answera 
to the animal individual, but only wliich 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.^ But, in botanical de- 
scription and classification, bj* the individual is meant the herb, 
shrub, or tree, unless otherwise si^ecified. 

C2o. Species in biological natuml history- is a chain or series 
of oi*ganisms of which the links or comix)nent individuals are 
parent and offspring. Objectively, a s^wcies is the totality of 
beings which have come from one stock, in virtue of that most 
general fact that likeness is transmitted from parent to progeny. 
Among the many defmitions, that of A. L. Jussieu is one of the 
briefest and best, since it expresses the fundamental conception 
of a species, t. e. the perennial succession of similar individuals 
peqx»tuated by generation. 

C)'2G, The two elements of species are : 1 , community of origin ; 
and, 2, similarity of the comiX)nent individuals. But the degree 
of similarity is variable, and the fact of- genetic relationship can 
seldom be established by obser^'ation or historical evidence. It 
is from the likeness that the naturaUst ordinaril}' decides that 
such and such individuals Iwlong 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 branch or stem, the phytomers of the branch are repetitions and 
progeny eacli of the preceiling one, so forming a series of vegetative 
generations ; and the whole tree might almost as well represent tlie 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. Ix^ast tenable of all is the conception that the whole product 
of a seed may lie taken to represent the vegetable individual. For then 
individuals incrcasi^'d by buds and division are wholly unlimited Imth in ex- 
tent and in duration, so far as obsi'n'ation can show, and a multitudinous 
race, not only of the pn'sent and past, but perhaps in jierpetuity, may con- 
sist of a single individual. There are, indwd, theoretical reasons for infer- 
ring that a bud-propagated race may not last so long as a seed-propagated 
species ; but there ii no proof of it. See Darwiniana, Art. xii. 



G27. No two individuals are exactly alike ; and oflbpi _ 
the same stftek rofty lUffei- (or in Ihtiir progeny may coir.e to dilferjfl 
strikingly in some |)articulars. rSo Iwo or more forms ■ 
would liave been regarded as wholly distinct are aomctiiiu 
proved to be of one species by evidence of their common oiigio j 
or more conimonlj' are inferred to be so from the observattou c 
a series of intermediate forms which bridge over the ditlei-enct 
Only obsen'ation can inform ns bow mncli difference is comnt 
ibic with a common origin. The gcncml result of observatioi 
is that ])lants and animals lireed true from generation to gener»-f 
tion within certain somewhat indetei-rainatc limita of variatioii ;j 
that those individuals which resemble each other witliin end 
limits interbreed (Veely, while those with wider diflerences daM 
not. Hence, on tlic one band, the naturalist i-ecognizes Varietiet ' 
or differences wiHiiii the si>ecies. and on the other Genera and 
other 8ni)erior associations, indicative of remoter relationship of 
the species themselves. 

G28. Y'arlctiea are foi-ma of species marked by characters of J 
less ftxity or im|iortancc than arc the species themselves. Thejrl 
may be of all grades of difference fixmi the slightest to the mosfe* 
notable: they abound in free nature, but assume partivv 
imiwrtancc imder domestication and cultivation; under nhict 
^'aliation8 are most pi-one to oiiginale, and desirable ones i 
presened, led on to further development, and relatively flxed. 

629. If two eeeils (Vorn tlie same ikkI are sown in different^ 
soils, and submittei:! to different conditions as re8i>ecta heat, li^t, I 
and moisture, the plants that spring from them will show morkl 
of this different treatment Iji their appearance. Such dilfbrenoefl^ 
arc continually arising in the natural coui-se of (liings. and toM 
pi'odnce and increase tliem artificially is one of the olijwU rf.W 
cultivation. Striking as they often are (esjiecinll.i' in anitiuls ] 
and biennials), they are of small scientific consequence. Wbea> j 
spontaneous they arc transient, the plant cither outlnsting ths' 
modifjing cause or else succumbing to its continued and graver 
oifcration. But, iu tlie more marked varieties which alone de- 
ser^-c the name, tlie cause is occult and constilulioual ( the 
deviation occurs wc know not why. and c'onliuucs throughout 
the existence and growth of the herb, shrub, or tree, and con* J 
seqnently through all that proceeds IVom it by propagation froial 
buds, as by offsets, layers, cuttings, gratis, &c. 

G30. Some varieties of cultivation originate in comparative 
slight deviations Ttvm the t\-pe. and arc IhI on to greater differ-l 
enees by strict st^^lcction of tlie most marked individuals lO;! 
breed l>om. Most appear as it were full-tledgcd, except as tO'J 


luxuriance or development, more or less under the control of 
conditions, their origin being whoUj' unaccountable. They arise 
in the seed-bed, or sometimes from buds, which as the gardeners 
say *' sport'* ^ That is, some seedlings, or some shoots, are 
unlike the rest in certain particulars.* 

G31. 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 *' sporting" of certain buds 
into characters in branch, flower, or fruit unlike those of the 
stock, is known in a good number of plants.' It might also 
oocur in corals, hydras, and other comix)und animals propagated 
by budding. Once originateil, these varieties mostly persist, 
like seedling varieties, through all the generations of budding 
growth, but are not transmitted to the seed. 

G32. Upon the general principle that progeny inherits or tends 
to inherit the whole character of the parent, all varieties must 
have a tendenc}* to be reproduced by seed. But the inheritance 
of the new features of the immediate parent will commonl}' be 
overborne by atavism, t. e, the tendencj' to inherit from grand- 
parents, great-grand-parents, &c. Atavism, acting through a long 
line of ancestr)*, is generally more powerful than the heredity 
of a single generation. But when the offspring does inherit the 
peculiarities of the immedis^te parent, or a part of them, its off- 
spring has a redoubled tendenc}' 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 oveqx)wer the influence of remoter ancestr}'. Close-breed- 
ing (398) is requisite to this result. In the natural wild state, 
varieties — man}' and conspicuous as they often are — must be 
much repressed b}- the prevalent cross- fertilization which takes 
place among the individuals of almost all species. Cultivators and 
i)rceder8 in fixing varieties are careful to secure close breeding 
as far as this is possible. This has fixed the particular sorts of 
Indian Com, Rj-e, Cabbage, Lettuce, Radishes, Peas, &c., and 

1 Both the technical English term, Sport, and its Latin equivalent, TMaus, 
arc 8omctini(.^ used for bud-variation only, yet as commonly for seedling 
variation also. 

3 l>arwin 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. Naegeli, Braun, 
and myself incline to the opinion that each plant has an inherent tendency 
to variation in certain general directions. 

* A list of known bud-varieties is given in Darwio'i Variation of Animals 
and Plants under Domestication, Chapter xL 



indeed of nearly all our varieties of cultivated annual and liie 
esculent plants, as well as of several perennials, many of which 1 
have boon fixed through centuries of domestication, while others [ 
are ot rticcnt ostAbliFhineut. What ia now taking |)Iaee with 1 
the I'each in this country may convince us that lieritablc varieties J 
may lie develoi)eil in trees as well as in herbs, am) in the same \ 
manner; and that the reason why most races are annuals or 
biennials is because these can be periictuated in no other waj, 
and because the desired result is obtainable in fewer years than I 
in shrubs ur trees. Varieties of this fixity of character are called J 

G33. Bsces (Lat. Proltt). A race, in this technical » 
the term, is a variety which is peipetuate<l with oousidcrablii j 
certainty by sexual propagation. This distinction of varieties 
pertains chiefly to I>otanj-. In the animal kingdom all penuanent I 
varieties must be races. So are all indigenous varieties of 
plants.' In most of these, the [Hwition of siwoies and variety is | 
more or leaa arbitrary or accidental, and capable of interchange. 
What is called the siM^ciea may be only a commoner or betler- 
known form, or the one first recognized aud nanieil by botanists ; 
whence the other forms as they come to be recognized arc made ■ 
to rank in the books as varieties. Instead of one varying ftom i 
the other, all the forms have probably varied ages ago fVom k 
common tyjK. 

G34. These varieties of the higliest order and most marked | 
characteristics, licing peqwtuable bj' seed, have the principtU | 
attributes of species. They are a kind of suboixlinatc derivative \ 
species. Hence they are sometimes calletl Subspfciea. We J 
judge them not to be so many species, either because in the c 
of cultivated races we know something of their origin or historj*, j 
and more of the grave changes which long domestication may I 
bring to pass; or l)ecause the forms, however stable, H{Ba | 
among themselves less than recognized species generally do ; or I 
hecjtuse very striking dilfeiences in the extremes are connected [ 
by intermefliaU- forms. And our conclusions, it must \m- iindsr- j 
stood, ■' are not facts, but judgments, and lai^ely fallible Judg- I 
ments." ' For while some varieties ap|>ear strikingly dilTenDtt t 
some species are very much alike.* 

' Till' llortcrailisli luiil a few nihcr plaau of «iKinlancous itmwtli. which 
ihrouBli long <lcp#iii1en<.-e on bud-prupatcatlon >««in lo hare losi Ihe power 
of •clring tM'il, oar hsnllv be caIImI vnrielice. 

1 DarwinUti*. .tJ>, 

' \T)H'n-run'. tinw wr hintl.r need the term mx in the miricled WOH 
of wpd-pn>p>|i>t«l raririj'. it i^ mmctiniei (.'mivrnltrnl to luc il Id Ihe nwn> 
uvr prupuK'd by UcDt1i4ni (Anniveriaiy AJdrcu to tlie Liuncaa Socletji, 


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 freel}' 
as do those of the same variety and are equally prolific. Their 
union produces 

63G. Cross-breeds.^ In nature, cross-breeding doubtless re- 
presses variation or prevents the segi'egation of varieties into 
what would be ranked as species. In cultivation and domesti- 
cation, it is tume<l 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 indi>iduals of distinct species do not 
interbreed, although many arc capable of it. There is great 
diversity* in this regard among plants, some (such as Willows, 
Verbascums, and Verbenas) interbreeding freely and reciprocallj* ; 
some interbreeding in one direction, but not reciprocally ; others, 
even when ver3' similar, refusing to unite. But, on the whole, 
there seems to be few nearlj' related species in which the pollen 
of the one cannot be made to act upon the ovules of the other 
by peraistent 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 pla}' a very prominent part in nature, apart from 
cultivation, although the limits of some species may be obscured 
by them, ix)ssibl3' of more than is generally supix)sed. In the 
animal kingdom, all the most familiar h^'brids are sterile : in the 
vegetable kingdom, a majority ma}' have a certain but very low 
degree of fertilit}* ; but this is also the case in many unions 

May, 1800, 5) as the common designation of any group or collection of indi- 
viduals whose characters arc 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 i)lace 
as the designation of the highest grade of variety. The objection to this is 
that the subspcK^ific and specific names would be more liable to be confused. 
^ Half-breed is a common equiyalent term in the animal kingdom : Latin, 
MUtus or Mixtus ; French, MAis, 



within the Bpeciea, and espwially in the applicBtion of the pollen J 
to the stigma of the same blossom. Coranionly the Bterility of m 
bybiida is owing to the iinpotence of the staniens, which perfect 1 
no pollen ; and moat such hybrids may be feitilizc-d by the gmllea J 
of the one or tJie other parent. Then the offspring either in J 
the first or seeontl generation reveits lo the fertilizing species. 
Moreover, certain hybrids, such as those of Datura, which i 
flilly fertile ptr »e, divide in the offspring, partly in the first gen- • 
oration, and completely in two or three Buccecding genemtio&B, 
Into the two eomfxinent species, even when (.-lose-fertiliacd.' (In I 
part. this maycomc from adventive emhryo-forraation, 333.) 

639, Tliere appears, therefore, to be a real ground in nature 1 
for species, notwithstanding tlio difficulty and even impossibility I 
in many cases of defining and limiting them. 

G40. Spfciei is talicn as the unit In zoological and l>otanical 1 
classification. Important as varieties are in some respects, 
especially under domestication and cultivation, tlicy figure in 1 
scientific arrangement only as ll-actions of siwcies. S]>ecte9 
are the true subjects of classification. The aim of sjstematio 
natural history is to express their relationship to each other. 

641. The whole ground in nature for tlie classification of 8i>e- 
cies is the obvious fact that eiKcies resemble or differ from each 
other unequally and in extremely various degrees. If this were 
not so, if related species ililfered one from another by a constant 
quantity, so tlmt, when arranged acconling to tlieir rcsomblaiices, J 
the first diffei-ed from the second about as mueli as tlie second i 
fKim the thiiil. and the third from tlie fourth, and so on, — or if j 
the species blended as do the colors of the rainbow, — then, with J 
all the diversity in the vegetable kingdom there actually is. tlicrfrl 
conld be no natural foundation for their classification. Tlie tniil' T 
titnde of species wonlil render it necessarj' to classifv them, but i 
the classification would lie wholly artificial and arbitrar>*. Th« 
actual constitution of the vegetoble kingdom, however, as ap- 
{wars from observation, is that some sjiecies resemble each otlter 
very closely indeed, others differ as widely as possible, and be- 
tween these the most numerous and the most various grades of 

> Acconling lo Nauilin In ComplM R>-n<1ui, xUx. 1S5I>, & It. 1863. 8cs 
rIhi Nnudln's memoir nn liy brfillly In pl«nt> in Ann., Sci. KmI. bcT- 4, idx. 
ISaa, pp, 180-20.1, & in Mrni. Acail. Scl. . . . For tlie lilrralurf on TPgeUUa< J 
lijrbricli, ICC Kd-lreutpr, Nachrirhl. &u.. 1701, and Aptwndlcn. 1703-1708;. 
lIvrboM.on Amiryllitlaces. |Wi7 ; C. F. risnncr, Vpniiclii- iin<l Brolmchlnn-.] 
Ken urbpT die Bi»l«r<]iTEcuguiiK in PfliinienrciL-Ii, 1^11; Wiuhura, 1 
Butnrdbrfruclilung iin Flliinn'nrvkli. crliiutcrlcrt «u dun Butanlen iiit% 
WMden ; uiil the itHMiioir ot NauUln referred to. 


resemblance or difference are presented, but alwa3's with a mani- 
fest tendency' to comix)se 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 vanous associations the naturalist 
endeavors to express, as far as is necessary or practicable, by a 
series of generalizations, the lower or particular included in the 
higher 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. Linnseus 
and the naturalists of his day used names for only three grades 
of association, or groups sui^erior to species^ viz, the Genus, the 
Order, and the Class ; and these are still the principal members 
of classification. 

642. Genera (plural of Gentts) are the more particular or 
si)e<;ial groups of related species. The}' are groups of species 
which are much alike in all or most resi)ects, — which are con- 
structed, so to sa}', u[x>n the same particular model, with only 
circumstantial differences in the details. They are not neces- 
sarily nor genei-allj- the lowest definable groups of sixKxies, 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 man}' genera are so strongly marked, 
at least so far as ordinarj' obsen-ation 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 any other plants. The 
number of species in such a group is immaterial, and in fact is 
\QT\ 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 onl}' one species of Oak were known, the 
Oak genus would have lH?en as explicitly discerned as it is now 
that the species amount to three hundred ; and better define<l, 
for now there are forms quite intermediate between Oak and 



Chestnut. Fdmiliai' illustrations of genera In tiic anunal kiS^' 
flom are rumiMlieil hy the Cat kind, to which belong lliu domv^tjo 
Cat, the Catamount, the Piinllii-r, the Lion, the Tiger, the Lvo[i- 
anl, Ac. ; and by the Dog kind, whivh includes with the Dog 
tlie different species of Foxes and Wolves, the Jaeknl, &c. The 
languages of tlio most boi'lHirous as well as of civilized peo|ile 
everj-where show that they have i-ecognized such groui>8. Xatu- 
rnliftts merely give to tliem a greater degree of precision, and 
indicate what tlie [xiiats of agreement are. 

G43. If most genera were ns conspicuously marked as those 
from wliich these illustrations arc taken, genus would be as defi- 
nitely grounded in nature as sjicdes. Cut jiopularly recognized 
genera, rightly based, are comparatively few. I'opulai- nomen- 
claturc, ciuliodying the common ideas of people, merely sbows 
that generic grou|)s are recognizable in a considerable nnmbor 
of cases, but not that the whole vegetable or the whole animal 
kingdom is divisible into a dcHiiite number of such groups of 
equally or somewhat equally related species. The naturalist 
discerns the ground of genera in cliaraeters which the casual an<) 
ordinarj- observer overlooks; and, taking the idea of geiicnt 
trova the numerous well-marked instances as the nonn, applies 
It aa well as |x>s8ible to the less obvious or less nalural cases, and 
groups all known siM^cies under genera. Rcsemblanc-cs among 
the species when rightly grou]>ed into genera, though real, are 
often 80 unequal in degree, that certain species may be about as 
nearly related to neighboring genera. So that tlic recognition of 
genera even more tlmn of species is a matter of jn<lgment, and 
even of conventional agreement as to how and where a certain 
genus shall be limile<l, and wliat particular association of 8|)ec-lefl 
shall hold the position of genus. All the si»ecte» of a genus must 
aucord in everj- ini{>ortant structure ; but extended ohBer\'ation 
only can settle the question as to what are imiwrlant and what 
are incidental characters. For example. Ihe pinnalifid or slna- 
ftt« 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 AV'illow or Laurel. An open acorn* 
cup beaet with imbricate<] scales is a character common to all 
European ami American Oaks i but in numerous Asiatic species 
the cup bears concentric or spii-al lamellie instead, and in others 
the cup takes Uie form of a nak»l and closed sac. Maples have 
pal ma tely> veined and lot)e(1 leaves; but one s|M?cies has undi- 
vided and pinnately-veined leaves. The Apple and Uie Pear 
under one view are of the same genus, under another they rep- 
resent different gener^. 


644. The genus must be based on close relationship of species, 
but not necessarily on the closest. Raspberries differ from 
Blaekbenies, but must be ranked in the same genus ; and so of 
Plums and Chemes. For the groups which are to bear the 
gcneiic name must be as distinct and definite as possible. 

645. Orders arc to genera what genera are to species. The}* 
arc groups of a higher rank and wider comprehension, expressive 
of more general resemblances, or, in other language, of remoter 
relationship. As all s[jecies 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 zoolog}') have alwa3*s in botany been inter- 
changeable terms, and will probably so continue.^ 

646. As examples of orders in the vegetable kingdom take 
the Oak family, com|)osed of Oaks, Chestnuts, Beeches, &c. ; 
the Pine family, of Pines, Spruces, Larches, Cedars, Araucana, 
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 owlers 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. Mineralog}', Chemistr}', 
&c., ma}' use the same terms (genus, species, &e.) 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, Clou, Kingdom; or, in descending 
from the universal to the particular, 




1 Order is the older term, and that which associates best with the technical 
Latin names. Family is a happy term, which nssociates itself well with 
English names. But its use is attended with this incongruity, that the tnlte 
((K>$) in natural history classification is subordinate to the family. In 
zoology, order is distinguished from family as the next higher grade. 


650. This is the common IVamcv^ork of nattiml history cR 
ficntion. AH plants and all aniuiaU belong to some specit- 
everj" siwcies to some genus ; every genua to some onlcr or J 
familj- ; everj- order to some class ; every class to one or the J 
other kingdom.^ But this framefrork, althongli all that is K^- J 
qutsite in some parts of natural hiatoiy, does uol express all the 1 
obserrable gradations of relationship among si»ocies. And vwu J 
gradations l>e]ow species have sumi'times to lie classified. The I 
series is capable of extension ; and extension is oflcn requisite I 
on account of the lai'ge number of objects to be arranged, and ] 
tlie various degrees of relationship which may come into view. 

651. This is efTecIed by the intercalation of intermediate 1 
gi-odes, to he introduced into the system only wlien there is 1 
occasion for them. And in botany one or more grades superior J 
t« the classes are neeilfiil; for first and foremost is the great I 
division of all plants into a higher and a lower Series ' (or aub- [ 
kingdom), the I*hKnogamou8 and the Crv])togamons. 

6a2. The grades intercalatctl into the long- established sequence I 
of Class. Order, Genus, and Species, with new names, are luaiiily 
two. Tribe and Cohort. 

053, Tribe has been for a generation or two thoroughly estab- | 
lishe<1 in both kingdoms, as a grade inferior to order and supc- j 
nor to genus. In botanical classification, much use is iiuulc of | 
tliis grade, genera lieing grouped into tribes. 

6i>4. CVthort (Lat. Cohora) is of more recent introduction, at | 
least in Uotany, but is Itccoming established for a grade next \ 
above that of order. Oi-ders are grou|je<l into cohorts. Undley I 
hit upon a goo<l English name for this grade, tliat of Allianet. 
But this wortl has no a\'ailublc Latin equivalent ; while ct^ort I 
takes equally well a Lntin or an English Ibrm. 

G55. Finally, each grade is capable of l>cing doubled by tbe I 
recognition of one like it antl immediately subordinate to it. aaH I 
with designation directly expressive of the snborrlinatlon. For | 


' Aniwcring to the Frpnch Etahnuiehrmeal in tmlngy. For lhl» ft i* 
pnspil In vtf the word Oiciuon {DivUio] : see Lawi of Dainnlcal Kamencli^ 1 
turc siloplt^ by tlie Inh-malinnRl Botanical Congrme held at I'ari* In I 
Auinut.lMT: loft^thpnviih&ItistorlL'allntmtuoionanilaCiiniinpniarjr.tqr I 
Alph. Drranilollp, — F.n|[llBli tranalalion, Lonilun. I»I8: IheoriitlnalFmich I 
(tdilloa. Paris, 1807. Perhipe no Viler name can be tnunil ; 1ml the ~ 
DeCaiidollv lirnu^hl Diviiio iniQ cominon hm fur a gni\i- ■iilxirilinatit tn J 
tribe. EixlUchcr ctuploj'cd ihe tenn Hegio. Wg have tiwU Utria, m 
mnch prefer it. 


example, if Dicotyledones and Monocotjledones be the two 
classes of PhaeDOgamia, the former (and only the foimcr) is 
divided uix)n very important characters into two branches, of far 
higher rank than the cohorts, viz. the Angiospermae and the 
GymnospermoB, which take the name of Subclasses. Ordens, 
esi^ecially the more comprehensive ones, often comprise two or 
mora gix)ups 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 gi*oups 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 im[>ortance and pretension (which maj- come to 
be allowed), tlie}' are tei-med Subgenera. Finall}', forms which 
are rankcil as vaiieties, but which may establish a claim to be 
distinct species, are sometimes termed Subspecies. Even what 
we regard as a variet}' may comprise more or less divergent 
forms, to be distinguished as Subvarieties. 

G5G. Some of the lai'ger and most diversified orders, tribes, 
genera, or species may require all these anal3'tical 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 : — 


Series or Diyision, or Sub-kingdom, 



Order or Family, 



Subspecies or Race, 


6.57. Katare 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 accorcl only in the most general respecte, in that 
which makes them plants rather than animals. Plants of the 
same variety are generally as much alikp as if they were of the 


BAme ioimediatc parentiige. All plants of the same Bpede^^^ 
so ranch alike tliat tliej' ate iDlcrn^tl to have deucended fVom 
a commuu stock, aud tlifir difTcrent-es, iiowt'ver grave, are sup- 
posed to have arisen ft'om subscqiieut variation, and llie more 
marked dillbrences to have Itecoiue fixed tlirougli lieredity. TItis 
is included in ibc idea of species. Desueiit n'om a commoa 
origin (.'xplains the likeness, and is the only explanation of it. 

6j8. But what is the explanation of the likeness between the 
species themselves? Aa respects nearlyrelated s|iecies, the answer 
is clear. Except for piactieal pnr[x>se8 ami in an arbitrary 
way, no certain and unfailing distinction can be drawn between 
vant>ties of the highest gnule and s|)ecies of closest resemblance. 
It cannot reasonably be doubted that tliey are of simUar origi- 
nation. Then there are all grailationa between very closely aud 
less closely related species of the same genus of plants. 

G39. The Theory of Oesreot, that is, of the diversilicaliou of the 
species of a genus through variation in the lapse of time, utfords 
the only natural explanation of their likeness which has yet been 
conceived. The alternative snpitosition, that all the exisiiiig 
species and forms were originally createtl as they are, and luve 
i-ome down essentially unchanged from the li^inning. offers no 
exphinatiun of tlic likeness, anO even assumes that there is do 
seientitle explanutiun of it. The hypothesis that the si^eciee of 
a genus have become what they arc hj- divei-sification through 
variation ia a very old one in botany, and has from time to Umo 
been put forwaiil. Hut until recently it has had little influence 
ujwn Uie science, Iwcause no dear idea had been fonned 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 eiiually 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 diffur- 
e»t as those which we recognize as species. But this would not 
account for the limitation of species, which is the usual (bat 
not universal) characteristic, and is an essential part of the 
idea of siK-cies. Just this is accounted for by 

6G0. Natnrol Selection. This now familiar term, proixtsed by 
Charles Darwin, was suggested by the operations of breeders in 
tlie development and fixation of races for man's use or fiincy;— ~ 
In aniTnals by breeding from selected parents, and selecrting for 
breeding in each generation Uiosc individuals only in which the 
desired points are appaiTUt and predominant ; in tlic aeed-betl 
by rigidly <leHtro,\"ing all plants which do not show some desirable 
variation, breeding in and in fi-om these, with strict selection of 


the most variant .form in the particular line or lines, until it be- 
comes fixed by heredit}- and as different from the primal stock 
as the conditions of the case allow. In nature, the analogous 
selection, through innumerable generations, of the exceedhigly 
small i>ercentage of individuals (as ova or seeds) which ordi- 
narily are to sur^•ive 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 
exi>osed. In the Struggle for Life to which they arc thus inevi- 
tabl}' cxjjosed, only the individuals best adapted to the circum- 
stances can survive to matuiit}' and propagate their like. This 
Survival of the Fittest^ mctaphoricall}* expressed b}' the phrase 
natural selection, is in fact the desti'uction 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 thereby adapted to different conditions, some to one 
kind of soil or exi)osurc, some to another, thus lessening the com- 
petition between the two most divergent fonns, and favoring their 
preservation and farther separation, while the intermediate forms 
perish. Thus an ancestral t3'pe 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 \y\Hi^ 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, 
geiUTa are later branches, species the latest definitel}' developed 
ramifications, varieties the developing buds.* 

GGl. Except as to those changes in size, luxuriance, or depau- 
l)eration and the like, in which plants, especially seedlings, 
resix>nd promptly to external influences, as to heat or cold, 

^ 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 tlie 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 Cliarles Darwin, also On 
the Tendency of Varietit»8 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 doctrine, see Darwin's " Origin of Species by Means of 
Natural Selection," " Tlie Variation of Animals and Plants under Domes- 
tication," and various other works; Wallace's "Geographical Distribution 
of Animals/' &€. For some ex|K)eitioii8, see Gray's " Darwiniana." 


moisture or Arynesa (which are transient and coraparativclj' 
tiuiinjKirtant) , vai-iation, or the uiilikeiieas of progitny lo parcoti ' 
is ooi'ult and iitexplii;at)le. If sumetiincs tilled out b}' the I 
external conditioDB, it ia bj' way ol' internal iV8[>oiise to tbem. 
In Dorwin's conception, variation of iteolf does nut tend in any i 
one i>artiuular direction : he ai>i>ears to attrilmtc all adaptation [ 
to the sorting which results from the struggli> for existence and I 
the survival of the fittest. We have Btipposed.and Nujgeli takes ] 
a similar view,' that each plant baa an internal tendency or pre- | 
disi>ositiun to varj' in some directions rather than others; &oia | 
which, under natural setettiou, the actual dilfeivuliationa and j 
adaptations have proceeded. Under tliis assumption, and taken ^ 
as a working liypotliesis, the doctrine of the derivation of spcciea 
serves well for the co-oixlination of all the facts in botany, and 
atTords a prabable and reasonable answer to a long series of 
questions which without it arc totally unanswerable. It is sap- i 
jwrted by vegetable paheontology, which assures us that the J 
plants of the later geological peiiods are the ancestora of the j 
ataual flora of the world. In accordance with it we may explaiUi 
in a good degree, the present distribution of species and other 1 
groups over iLe world. It rationally connects the order of tlie j 
appearance of vegetable ty|)cs in time with the grades of diflti^ i 
entiation and complexity, l)oth prooecding f^um the simpler, or ' 
lower and more general, to the higher and more <Iilferentiat«d 
or special ; it exjilains by inheritance the existence of function- 
less parts ; throws light upon (he anomalies of parasitic plants in 
their various gradations, u})on the assumption of the most various 
Ainetions by morphologically identical organs, and indeed illumi- 
nates the whole fleld of morphology with which tliis volume hxa 
been occupied. It follows that sitecies are not " simple curiosities j 
of nature." to be tatalogue<l and described merely, but that they 1 
have a history, the recunls of which are impi^ssed upon thdr 
structure as well as traccahle in their geographical and paheon- 
tological distribution. This ^'iew, moreover, explains Uiv r^ 1 
markable fact tluit the characters in which the afiinitios of 
Ijlanta arc mainly discerned (imtl which therefore serve beat I 
for ordei's, tribes, and other princiiuil groups) ai'c cunmionty such I 
as are e\ i<leutly of small if any imix>rtance to the plant's ^ 
being, and that they run like threads tiu-ough a seiies of Species J 
of tlie greatest diversity in habit, mode of life, and i)articidirl 
adaptations to conditions." 

1 Entiichnngunil BcgrifFdernilurliUliirigclienArt. Zwplir Aoflagr, ISOGt. 1 

' Thi* i( a L-omllary of nutural (election, wlili-h can lake rffccl oBlf t 

upoD UBctul cWsL'ten, i. e. upon itructurt* wliivh pUy »unie at-live ( 


662. The fixity of species under this view is not absolute and 
universal, but relative. Not, however, that speciilc 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 tertiar}' 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 

Section II. Botanical Classification. 

663. Natural and Artificial Classiflcatioiis ma}* 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 difierent in their principal structure. Arti- 
ficial classifications single out one or more points of resemblanc*e 
or difference and arrange by those, without reference to other 
considerations, convenience and facility being the controlling 
principles. The alphal)etical arrangement of words in a dic- 
tionary, and the sexual s^'stem in botany by Liniueus (or rather 
a part of it), — in which plants are arranged in classes upon tlie 
niuul>er of their stamens, and in orders upon the number of 
pistils, — are examples of artificial classification. The arrange- 
ment of the wonls 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 tliercfore 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, Rubiacese are known by insignificant stipules, Ano- 
nacese by ruminated albumen, Rhamnaceae by a valvate calyx and stamens 
bi>forc 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 f/enealoffical or genetic on the other. The saga- 
cious naturaUst seizes upon the latter for orders and the like ; wliilc the 
former are prominent in genera, &c. 



CG4. Ko artificial clft8sifii;ation of plants conlcl fail tol 
nuttiral iti Bomc portions ami some res[>octs : bocause pbinttf 
ivliich agree in any (Joint of strueture likely lo ha ujhhI for tlie 
purpose will cumruoiiU' *gre« iu uUioi' antl |>erhaps more tinpor> 
tant eliaraet«rs. t)n tiiv oilier hand, no natimil claseifiiwtion (.fio 
(lisjioiitie wilh artificial hel|JS ; Dor can it express in lineal ortliFT. 
or in any otiif r way, all the various relationsliips of planta, vvou 
if tliesc were fully detcruii tied and ri^litly subonlinated. Xatu- 
raiiiita now en<leftvur to make elasiiifi<;ation as natural as possible ; 
Itiut is, to base it id every grade u|K)n real relationships. Wtiat 
real relationsliipH are, and how to exjiress them iu a general 
system and Ihrougliout its |iarts. lius been the task of Uie leaders 
in botany from tlie beginning of the science until now ; and the 
work is by no means completed. 

I^ij^. Linnaius was perliups the first botanist to distjngnisb 
clearly between a natural and an artificial classifieatioii. He 
lalH)red ineffectually u|>oii a natural class ideation of th« genera 
of plants into orders ; and ho devise<l an effective artificial clossi- 
flcation, which became so popular that it practically suiHrrsedod 
all others for mor« than half a century, and has left a |x-nuancnt 
impression u|x>n the s<.-icnce. The last generation of botatiista 
who were titiincd under it has not quite passed away. 

6C6. A]ite>LInniHn Classlflration. Litintcus, in his I'hilosopliia 
Butanica. divided systematiuts into betcroduK and ortliodox ; 
the former, those who classify* plants by their roots, heritage, time 
of flowering, place of growth, medical and economical uses, anil 
the like ; the Utter, by the organs of fructification. It is remark* 
able that all the orthodox or scientific classifications anterior to 
Liunieus made a primary division of the vegetable kingdom into 
Trees and Herbs, refening the larger slirubs to the fi>nn<'r an<l 
the under-shnibs to the latter. — an arrangement which Itegan 
with Theophrnstus and was eontinue<l hy Hay and Toumefoit. 

6G7, The tliree most important names in Iwtanicsd Uxonomy 
anterior to Linna-us arc those of Cesalpini, !tay, aii<l Tournefort. 
.Scienlilic botany commenced with the former, in ItuM'. in the 
latter half of tlie si xteentli century . Ho first used the cinbrjoand 
its cotyleilons in clnssifi»ition. distingmslie<I ditfereiices in the in- 
sertion of fioral parts, and. indeed (esccpting the primary divisioii 
into trees and beriw) , founded all princiiuil characters upon the 
organs of fYiict]fi<^tion, esjwcially njion the fhiit and seed. 
Conrad tiosncr of Zurich had sonn-what earlier ntcf^nized this 
principle, but Cesalpini first applied it. 

6fi«. A ccntnr>- later (l(iltO-90) this principle wns carried into 
practiL-e by Kivinus (a nnine latinized from Bachinann), of 


Leipsic, in a whoHy artificial classification founded on the corolla. 
His contemporar}' in England, Robert Morison, somewliat earlier 
began the publication of his great work, the Universal Ilistor}- 
of Plants. In this was first attempted a grouping of plants into 
what arc now called natural orders ; and these were defined, some- 
what loosely, some by their fruit, infiorescence, and flowers, othera 
by their stems, the nature of their juice, &c. But the two great 
systematists of the time, who together laid the foundations of 
modern scientific botan}', were John Ray in England and Joseph 
Pitton de Toumefort in France. 

0(59. 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 Dicotyle- 
donous and Monocotyledonous. These great classes he divided 
and sulxlivided, 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 nerveil 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. Toumefort'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 uix)n modifications of the corolla, and it overlooked the dis- 
tinction l)etween monocotyledonous and dicotyledonous embr3'os. 
Its great merit is that here genera, as we now understand them, 
are first established and defined, and all the sj^ecies then known 
referred to them ; so that Toumefort was justly said by Linmeus 
to be the founder of genera. Ray maj' be said to have indicated 
tlie primar)' classes, Jussieu (in the next centur}-) to have estab- 
lished natural orders, and Toumefort to have given to botany 
the first Genera Plantamra. 

671. Linntean Classification. Linnsus, the great reformer of 
botany in the eighteenth centurj', thoroughl}* revised the principles 
of classification, established genera and s|)ecies 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 imi)ortance in classification (probabh' 
without knowledge of the clear suggestion to this effect made 
by Burekhard in a letter to Leibnitz, printed in 1702) ; their 
functions, so long overlooked, being now ascertained. He also 



drew« tint and pnctical distiartiati between natnndand airtl*- 
Seul vInMiikatiotts (£63), ud definring all CT>deator$ to makt 
Uie ftmner aratlable, except for genera, be de\-iee<l a practical 
■ubatitntff for it, as a key to tbe gvnen, nz. ku niebnited 

C72. fleXMll Sfitcm, or amogement of tbe groera under Milt 
llcial ctasacB and onlefB, founded apoo tbc stanieos and pntUa. 
Altliough now oat of nee, this aftifietal classifivalioD baa been 00 
popular and inflaential, and baa left so deep an ini[HVAaion upon 
the scieocc ami espeoaUy upon tbc language of IxHanj. tbnt it 
neetU to t>e presented. The primaiy dirisions are Uie clasaes, 
twenty-four in number. But tbe 24th eUiss. Cn*i)tognmia, con- 
sists of planta wbi(.-h have not stamens and pistils and conse- 
quently no proper Rowers, antl is therefore tbe eounteri>art of the 
r(*m&ining twent'i'-three classes, to wbieb tbe corrcspouding name 
of [*hanerogamia or, in shorter form. Phienc^mia ( I*ha-u<^amous 
plants} has since been applied. These twenty-three classes are 
characterized iiy certain mudillcations and associations of tlio 
atamens, and have substantive names, of Greek derivation, ex- 
preasirc of their character. The first ele^-en comprise all plants 
with i»erfect (T. «. hermaphrodite) flowers, and with a definite 
number of equal and uneonnected stamens. They are distin- 
gnislied by the absolute number of these organs, and arc dee^ 
nated by names compounded of Greek numerals and the won] 
antlria (frotii Mrjj(>), which is used metaphorically fur stamcR, as 
follows : — 

CI«M i. M'tMtKnHiAini'InilcBallaac.-hplKntB'lilhoneiUaivnto tlieflower; 
■■ tn Hippurii. 
1. DuMUMrA, tlicMv with Iwo (Uineni, as In the Lilac. 
B. TaiANPau, wilh Ilircc iiamriu, at in the Vnli-riRn ind Iris. 
4. TirBikiiDKiji, with fiiur ■t>men«, ai in the SMbuiiu. 
6. PuNTAiiiiii)*, Willi Arc BraTiivna, tlie moit frequent caw, 
0. IlKxANbiiiJt, with *ix (lammi. m in the Ul.v Famil.v, Lc. 
T. IIbitamdmi*. itlth irvrn ilamens. ai in KnrHvhMiniii- 
8. OcTANoni A, wKh right ■Inmen*, a* In E-rening Prininwe anil Fnchda. 
(I. EHiRtxDniA. with nini' jtamr'm. m in the Rhuliarb. 
1(1. nKriHUHiA. with tc-n Maintti*, nt in Ithodtxtcnilmn and Kalmia. 
11- l>ni>KriNi>HiA, with Iwrlvc ■tamcni, a> in AMrum nnd tbe Mlipto- 
ncttp; cili-ndcd alio to int^luilc thoK with from tbirlL-cti lo nbw- 
Ircn itamrnR. 

C73. The two fliiececfling classes inchide plants wilh perfect 
flowers having twenty or more uneonnecte<1 stamens, which, in 

13. IcxWAMDMiA, art- InwncI on the calyx (prrJKj'miut}. ai in the Bote 

Family' ; anil in 
1& PoLtAxnaiA, on UwT»c«ptaole (hypogynous). at in ilw BuUmci^^ 



674. Their essential characters are not indicated by their 
names : the former merely denoting that the stamens are twent}' 
in number ; the latter, that they are numerous. — The two fol- 
lowing classes depend upon the relative length of the stamens, 
namel}' : — 

14. DiDYNAXiA, including those with two long and two short stamens, 

as in the majority of flowers with bilabiate corolla. 

15. Tetradynamia, those with four long and two short stamens, as in 

flowers with cruciferous corolla. 

675. These names signif}' 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. MoxADELPHiA (meaning a single fraternity), with the filaments 

united in a single set, tube, or column, as in the Mallow. 

17. DiADELPUiA (two fraternities), with the filaments united in two 

sets or parcels, as in Corydalis and in many Leguminosse. 

18. PoLYADELPuiA (many fraternities), with the filaments united in 

more than two sets or parcels, as in Hypericum. 

19. Syngekesia (from Greek words signifying to grow together), 

with the anthers united in a ring or tube, as in the Sunflower 
and all Compositse. 

676. The next class, iCs its name denotes, is founded on the 
union of the stamens to the st3'le : — 

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 indiridual ; as in 
the Oak and Chestnut. 

22. DifEciA (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 analc^ous 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 : — 











s d - 

Hrwa « 

dej^iovt^acdo-j^co «i?io 

A m3 ■■ ^ 

5 -. - 'A 


•^ ^ ^4^^ i»^ i»^ fM w^ 0) 

• M 




S3 -55 r • 


5 o 


B O 



ES S^is 

* ir s 

CCS • 
« X « . 

c £ z:^c 



ill 55 SJSI 

C c = £ 

V 4^ w i«% 

e 3 

oe - 

& 1. ^ V 

» 5 2 

>. >. >k s«» 


£ --3 

G > O 




r.^ ss 

£ = i 

C<5- « 








^ = u 
C Z t 



at — •- 






680. The orders, in the first thirteen classes of the Linnaean 
artificial system, deix?nd on the number of stales, or of the 
stigmas when the styles are wanting ; and are named by Greek 
numerals prefixed to the word ^ryni'a, used metaphorically for 
pistil, as follows : — 

Order 1. Moxogynia, those with one style or sessile stigma to the flower. 

2. DiGYNiA, those with two styles or sessile stigiuas. 

3. Trigynia, those with three styles. 

4. Tetragynia, those with four styles. 
6. Pentagyxia, those with fire styles. 

6. IIexagynia, those with six styles. 

7. IIeptagykia, those with seven styles. 

8. OcTOGYxiA, those with eight styles. 

0. Enneagynia, those with nine styles. 

10. Decagynia, those with ten styles. 

11. Dodecagyxia, those with eleven or twelve styles. 

12. PoLYGYNiA, those with more than twelve styles. 

681. The onlers of class 14, Didynamia, are only two and are 
foundetl on the pericarp, namely : — 

1. Gymxosperxia, meaning seeds naked, the achenia-like fruits of a 

4-parted pericarp having been taken for naked seeds. 

2. Angiospermia, with the seeds evidently hi a seed-vessel or peri- 

carp, /. e. the pericarp undivided. 

682. The 15th class, TctradjTiamia, is also divided into 
two orders, which are distinguished merely by the form of the 
po<l : — 

1. SiLicuLOSA ; the fruit a jilicle (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 dei>end merel}' 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 onlers of the 19th class, S3'ngenesia, are six, namely : 

1. PoLYGAXiA AQUALis, whcrc the flowers are in heads (the so^ialled 

compound flower), and all hermaphrodite. 

2. PoLYGAMiA supERFLUA, the samc as the last, except that the rays, 

or marginal flowers of the head, are pistillate only. 

3. PoLYGAMiA FRU8TRANEA, 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, whcrc cach flower of the head [or glom- 

erule] has its own proper involucre. 

6. MoNOGAMiA, where soliury flowers (that is, not united into a head) 

have united antliers, as in Lobelia. 



685. The 23d class, Pohgamia, has three orfersTi 
founded od the characters or the two preeeding rlnsses nml 
Iwaring llieir names, and the third named ajwiTThc same pnu- 
ciple, namely : — 

1. MoMzcijk, where boUi Kp«n>tcO soil perfect Oowi^rT bk found la 1 

2. DiiBCi*, where Ihey occupy two iliflerent plant: 

3. Tkii£cia. whcreone indiTiilual Iwan tlie pcrfpct, MDOlhrrltleitand*! 

nnie, and ■ tliinl (be piitlUale fluw«r«. 

686. The orders oT the 24th class. Cry]itoganiia, the Ftoirc|s.l 
less Plants, arc so man.v natural orders, and are uut definable f 
by a single character. They are : — 

1. FiLicKs. ihc Perot. 
S. Mitsci, the Mosaic 
3 Alga, whii'li, u Ee tt by linosiu, comprised tho Ucpntk'K. LiclicDi, .1 
&c.. >■ well ■« iho M.'Hwecds. 

4. FuNoi, Mtuliroaau, &u. 

687. In its day, this artificial system well Tulfillcd its purpose, J 
and was preferred to all others on the score of facility and deB>l 
niteiiess. Now no botanist would think of employing it, norT 
would it be clioscri for a key to genera, which was its only Icgiti- \ 
mate use. 

flK8. The Natural S;«teiii was nglilly appreciated by LiniuctH, J 
who pronounced it to be the first and last desideratum in aystfr J 
malic botany ; antl he early att«-mptcd to collocate most known 1 
genera under natural orders (e, g. Piperita^ Palmm. Scilamii*^^ 1 
Orchidr<r, Ainentttcf<f. Stc, sixty -seven in number, including his I 
four cryptogamic ortters) . but without definition or arrangement. | 
In Ills later years, he was unable to accomplish any thing mom. f 
The difficult problem was taken up by Linnirus's i-ontemporar] 
and corrcsiKtndent. Bernard de Jussleu. who plante<l the t>otan{e I 
ganicn at Trianon with plants Brnni>ed into natural orders, bat I 
published nothing. His pupil. Adanson. who when a younga 
man lived for several years in .Scni^al. and who was as mmark-f 
able for rctvntricity as for erudition and ability, [niblisbed in I 
1763. in his Families dea Plantes. the first complete M-Btem of I 
natural onlers. But he seems to have taken liltle fVom \ ' 
lC4ichcr, and with all his genius to have contributiHl little to the I 
advancement of the natural nyaleni. 

689. Anioine Laurent de Jussien. nephew of Bernard, bag 1 
been called (he founder of the natural sj'stem of iKilnny, ami to I 
him more Ihon to any other one person this honor may b« I 
ascribed. In his fienera riantanim secundum Onlinea Nalii-I 
, dlai>0iilii, 1789, n:ttural orders of plants, one liundr.-'d I 


in number, were first established and defined b}- proper char- 
acters, and nearl}' all known genera arranged under them. His 
primar}' division of the Vegetable kingdom was into Acotyledanes^ 
Mmocotyledone9^ and DicotyUdones^ adopted from Ray, with a 
change which was no improvement. For his Acotyle<lones, the 
Cryptogamia of Linnaeus, are the "plants without flowers" of 
Ray : they are, to be sure, destitute of cotyledons (though not in 
the manner of Cuscuta), because destitute of embrjo altogether. 
The Acotyle<lones forming his first class, Jussieu divided the 
Monocotyledones into three classes upon single arid artificial 
characters, namely ui>on the insertion of the stamens, whether 
hyi)Ogynou8, perigynous, or epig\nou8 ; and the Dicotyledones, 
into eleven classes on similar characters, preceded b}' a division 
into ApetalcB^ Afonopeta'a^ Polypetalce^ and Diclines irregulares^ i. e. 
first upon the character of the perianth, then upon the insertion 
of the stamens or in Monopetalse of the corolla. The following 
is the scheme : — 

Acotyledoncs Class I. 


Stamens hypogynous II. 

Monocotyledones \ perigynous III. 

epigynous IV. 

Dicotyledones . > 

Apetalous . . < perigynous VI 

Stamens epigynous V. 

>erigynous VI. 

>ypoffynous VII. 


' Corolla hypogynous VIII. 

perigynous IX. 

epigynous: anthers connate X. 
epigynous : anthers separate XI. 


Stamens epigynous XII. 

Polypetalous . \ hypogynous . . . . XIII. 

perigynous XIV. 

I>icIinous (also Apetalous) XV. 

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 lK»gan with the latter, the pha?nogamous or flowering plants, 
and with those having typically complete flowers.' On account 
of its convenience and the gn^ater facilities for studying the higher 
plants, this order has been commonh' followed ever since. His 
I»rimar^' division on anatomical structure, into Vascular and 
Cellular i>lants. was a backward step, confusing a i>ortion of the 
lower series with the higher ; and the duplicate names of Exo- 
gen<B and Endogena^ api)ended to Dicot3'ledone8e and Monoco- 


t;j'tedoDcre. &s it now npi^ears slioultl have been omitted. 

gnulos of tbc Candollean system superior to llie onicrs, ID their ] 

final fumi. are mninlj tbese : — 

Div. 1. VASCULAR (more properly PH^NOGAMOUS) PIJ.NTS. 
Cliis t. Dtcottledonocs or ExoognoI's. 

Subcloaa I. Thalamifloroi's : petals (dUtincl) "ml ''■■"'^ns on <he 

11. C'ALTciFLonoi-s^ pGUls (dUllnct or eonlegL-mt) And 
itamcns sdnate to the calyx. 

III, CoROLLiFLOROUB : peWls (mo«iiy (wnlcscent) not tA- 

nnle to ralyx, bearing Ilit »lnmens, 

IV. MoKOctlLAHTDEOcs: pctnli Wan ting- 

Clui It. MoNocoTVLEDOTiot'B Or ENDOOBNOca. (No subclsues.) 

DiT. IL CELLULAR (more properly CHYPT0GAM0U8) PLANTS. 

Class I. ^TnEooAMocs : nilh aeinat apparatiw, ami 
Vascular liiiue. lEi/nitrltKrit-FillrTt.) 
Only cclluUr lixsue. {itiitci and l/'poii«a.\ 

Ci.An It. AitriiiaAHOus ; dMiitutc of sexual or^ni and of other than J 
tellular IJMue. {Lichentt, Fang!, Alija ) 

691. Crvptc^amotis plants of all ordora are now known to Iw ^ 
provided wiib sesea; and the Jussiiean divisions of the Dico- 
tj'ledoiiea into Apefolee (including Dielines), Monopfta.'ir, and 
Poli/peialir, are generally [irefi-rrod to those of DeCaniloUe. Into 
tlie present views of tbc classification of ttic Cr^ptogauiia it ia 
unnecesaaiy here to enter. Their general nrraiigement itita 
classes. &c., ia not j'et well settled, and the whole laxonomjrl 
of the lower Cryptogams is at present in a state of transition. 

C9i. John Lindley in successive attempts (between 1830 ft 
1845) variously modified, and in some few respects improTedal 
the Candollean arrangement. But. as neither his groupings of tl 
natural orders nor tbc new classes which be adopted liave 
approved, his sihemes nce<l not be here presented. He mtist b 
creiliu-d, however, witli the first allempt to carry into cOtict i 
suggestion nuiile by Brown, tliat the orders should tbonuelvt 
be disposi'd ns far as possible into su|)erior and slrietly i 
groups. In Lindley's first attempt, such groujis of two g 
were proposetl, the lower called ni'xnt (tendencies), the lllf 
cohorti. In his later and largest work. The Vegetable Rio) 
dom. these were reiluceil to one, and the name of allisnM H 
ooinn). But this word lias no good Latin equivalent, Kod I 
tenn cohort (coAorj) is preferred. 


693. Robert Brown, next to Jussieu, did more than any other 
botanist for the proper establishment and correct characterization 
of natural ordera. Having in the 3'ear 1827 published his dis- 
covcrj* of the g3mnospcrm3- of Coniferae and C^cadacece, it 
was in LiiKlley's works that this was first turned to proper 
systematic account b}* dividing the class of Dicotyledones into 
two subclasses, the Angiosperma and the GymnospernuE. The 
latter has been elevated hy the vegetable palscontologists to the 
rank of a class. 

604. Stephen Ladislaus Endlicher, of Vienna, a contempo- 
rary of Lindlc}', of less botanical genius, but of great erudition 
and aptness for classification, brought out his complete Genera 
Plantarum secundum Oi-dines 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 primarj' division is into two 
regions: 1. Thallophyta^ plants Without projKjr axis of growth 
(develo[)ing upward as stem and downward as root), no other 
tissue than parenchyma, and (as was thouglit) no proper sexes. 
This answers to the lower or Amphigamous Cellular plants of 
DeCandolle. 2. Cormophyta^ plants with an axis (stem and root), 
with foUage, &c. The Cormoph3'ta, or plants of the higher 
region, Endlicher divided into three great sections: 1. Aero- 
hrya^ answering to the higher jEtheogamous Cr3'ptogamia of 
DeCandolle, with which was wrongl3' associated a group of root- 
parasitic fiowering plants (the Rhizantheo?) which were fancied 
to bear spores instead of embr3-o in their seed ; 2. Amphibrya^ 
which answer to Monocotyledones ; and, 3. Acramphihrya^ which 
answer to Dicotvledons. These last contain five cohorts: 1. 
Gymnospermea ; 2. Apetala ; 3. Gamopetala (the Monopetalse 
of Jussieu I Hotter named) ; 4. Dialffpetalm (the PolyiJetalae of 
Jussieu, &c.). The cohort in Endlicher's classification, it will 
l)e seen, is a higher grade than that to which this name was 
applied b3' LindleN* in the more recent use. For the latter, i. e. 
for the grade between these and the order, Endlicher emplo3'ed 
the name of class. 

695. Finally, the Genera Plantarum, now in course of pub- 
lication b3' George Bent ham 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 G3Tnnospennou8 ; the former into 
the Polvpetalous, Gamopetalous, and Apetalous di\'isions ; and 
the first of these into the Thalamiflorous, Disciflorous, and Cah'- 
ciflorous ^^ series " (the middle one composed of the latter part 


gf DeCandoUc's Tlulamiflone with some of his Calfdfione) | 
aii'l uiidiT these Itic onlcre arc arranged in uuhoilA, — finn-n I 
ouhorU In Die I'olj"|ie1ala!. and ten under lbrc« •' scriee " in lli3 I 
<'tamo\K-Uihv. 1*116 reniaitidiT uf this particular claKsifivaiion T 
ban not yi-l ninx-ared in print, altbough partly HkL'tohiKl by its J 
authors. It will gi-nL-raliy Ik- adopted in this euuntr}', with some 
ocranioiuil minor niodlDcationa. 

C'J6. VariouH Rtoditii-'atlouft tiavc been froni time to time prtf 
pow-d. One of the Iwdt of them in principle is that iDitiated lijr 
Adolphe Brongniart and adopted hy many European botanists, 
which, recognizing that most apetaloue dowers arc reductions or I 
degrailatioiiH of [Mlj'pctalous types, intercalates the Apetalv or I 
Moiiot-liluniydctc nnioug ilie Polyix-talte. Bui, this has never yt^ I 
been done in a Bathfoctoi^- manner, or without sundering ordcra 1 
which Hlioidd stand in i-ontig:uUy. 

CU7. It Hlionld )>c borne in mind that the natural system of J 
botany in notural only in the constitution of its genera, tribes^ J 
nrdent, &c., and in its grand divisions ; that its cohorts and tlie I 
like are as yi^t only tentative groupings; and that the puttingi 
tJ>gt-tbi-r of any or all these purls in a system, aud. especially' 
in n lineal order, necessary ns a lineal arrangement is, mnsk'l 
needs l)c largely artifiml. So that even the- best iterfected I 
armngements must always foil to give of themselves more tliani 
ail imperfect and coneiderabl}* distortcl reflection of the plan oT f 
the vegetable kingdom, or even of our knowledge of it.' 

■ In the flnt pUn. thi- rclationnliips of any group uinnot always I 
rlKhlly ollmatcti bvforc all ill monibcn are known anj tbdr whole iirwrt J 
mv unJi'rdfKxI ; >o that ihe viewi i>f bolaniaii arv liable b) be niodifled wftkl 
ihu illw-uverli'i of over> year. Tli« diw.tiver)' uf a ningie plant, a 
u( (triiuturtr iK-tun? miiundi^ritoodi luut >umeliiDt.-i cliannpri] malcriatljr tl 
p«ltl(m of B conililfniblc group in Ihc lyilcra. and minor alicraiiuni ai 
cunllnually maitr by our iniTeaiing knowledge. I'hm Ihc groupi whiL-h « 
T(vo|[nlio, and diailriguiah *• genera, trib«i. ord^n, &c.. >rv nui alwaja, oj 
[wrliap* ncii (tcnprally, coniplvU'lyeircinnacrilivd In nsliirr. a> wcar« oblig* 
Id ajiunie tht-m lo hv In our uUiilflt-aliun. Tliii inielil be vxpwied fro 
the nitnrc of the inuc. For iIig naturalUt'* groupt, of whalcTcr gnd*. ■ 
niil rralitin. but idta: Tlirlr eoiuidenitlon involve) quenlinns, not of >!' 
Iirlv«i>ii which abiolute dittlni-liona might l>e drawn, bill of dryttn of n 
UiiHrr, wlili'h may be expi-c'tcd to pmcnt inftnJtu gmdaii'iiiK. llv*id««, ■] 
though tlic Krailca nl alflnily amunB tppclea atv must rariuui, If not wboDj 
initpllnile, Ihv iiaturallat rrdu(.-« them all to a t>'W, and trvali hit gtaiHa 
iribu, kv., ■> i-qoal unit*, or a* dlitinguiahed by oharaclen of about rqM 
value lliroughout, — whieb i* far (mm lieliig ibe caae. And in hii wo ~ 
Iw ii obll|ti'd lo amniit- Ihe urollpa \w n-eiiKn'KCB in ■ liiii-al aeriM; '■ 
uai-li igfmut or orliT. fte . ia *i-rj often about eiiunlly ndnipd lo three or t 
ollien ; an thai only a part of (lie telallonahlp of plants can In any w*; |i 
Indlealtd by a Un- •niluent. 


G98. Even the great classes cannot be arranged in a single 
line, beginning with the highest Phflenogams and leaving the 
lowest in contiguit}' with the higher Cr3i)togams. The Dieot}'- 
ledons take preceilence of the Monocotyleclons in rank. Yet a 
part of them, the Gymnospenns, are much the lowest of all 
known Pha;nogams as reganls simplicity of floral structure; 
and through them only is a connection with the higher Cryp- 
togams to be traced. The Monocotyledons stand u^wn 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 tendency, we raise the Gymnosperms to the 
rank of a class, and place it between the Monocotyledons and 
the Acrogens, then the much nearer relationship of G3'mno- 
sperms to Angiosperms through Gnetaceae and Loranthaceae is 
not respected. (60G, &c.) 

699. Nor can the angiospermous Dicotyledons be disposed 
lineally according to rank. The apetalous and achlamydeous 
must be the lowest. Some are evidently reduced forms of Poly- 
petalae or even of Gamoi^etalae : the greater part cannot without 
violence be thrust into their ranks. The Gamopetalie, esixjcially 
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 Compositas 
l)erhaps at their head. In the CandoUean 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 homolog}* 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 
alK)rtions 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 studie<l in 
the S3'stematic 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 stnicture, relations, and most important products of the 
plisenogamouB natural orders, as another is to illustrato the 

844 TAX0N03IY. 

cryptogamouQ orders. A 8}'noptical A'iew 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 cliapters. 

sekip:s l pilenogamous or flowering plants. 


Slbc'lahs I. AXGIOSPERMS. 


I^ic. 2. Gamopetalol'8. (Monopetalous.) 
iJiv 3. Apetalous. 


hiv. 1. Spadiceous. 
hiv. 2. Petaloideous. 
Die. S. Glumaceol'S. 


Class 111. ACR()0P:NS. 

iJic 1. Vasctlar. (Ferns and their allies.) 
Die. 2. Cellular. (Mosses and Liverworts.) 





Section I. Nomenclature. 

701 . Phttography 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 Glossolofft/ or (to use the more 
common but less proi^er word) Terminology, The latter is prop- 
erly Nomenclature. 

702. Names of Plants were at first onl}' generic names. The 
language of botan}' 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. Quereus, Prunus^ Rosa^ Bubus, 
Trifolium ; and of Latinized Greek names, Agroitis^ Aristdochia^ 
Colchicum^ Melilotus. To the classical names others were 
added from time to time ; as, from the Latin, Bidens^ ConvaUarta^ 
Dentaria ; from the Greek, Anacardium^ Glycyrrkiza^ Lomnthus^ 
&c. Some barbarous or outlandish names were early adopted, 
such as Alhagi from the Arabs, and Adhatoda and Nelumho from 
India. These are mostl3' such as were or could be conformed to 
Latin ; as Datura and Ribes from the Arabic, and later l^hcea and 
Coffaa. Of American aboriginal names, Hara^ Guaiacum^ and 
Yucca are examples. Some ancient names of plants commem- 
orated distinguished men. Ex. Ascleptas^ Euphorbia^ Lysimachia^ 
PtBonia. TouiTiefort and his contemi>oraries resumed this 
practice, and named plants in memory or in honor of distin- 
guished botanists. Ex. Begonia^ Bignonia, Caescdpinia^ Fuchsia^ 
Gerardin^ Lobelia^ Lonicera^ Magnolia. 

703. When among plants of the same name or kind diff*erent 
species were known, these were distinguished by annexe<l epi- 
thets. For example, among the Pines there were : Pinus syU 
restris, vulgaris ; Pinus sylvestris^ morUana altera ; Pinus sylvestris^ 
montana tertia ; Pinus sylvestris maritima^ conis firmiter ramis 
adfuBrentibus ; Pinus maritima minor ; Pinus maritima altera^ &C« 


Ami as the number of known s]M>cies increaaeiJ. ' 
biiglb of tlie iilirasc'S wliit-U were needed for thi'ir cliscritniDa- 
tion, TlicBu ■" clitlii-eiitiip," tliiis used as six-dfio nnmcs (the 
iiumina t/iecifica of Liniiieus), became extremely cumbroua. It 
was about iu the imd<lle of bis career that Linnieus suggested 
wbat lie calltHl trivial names (rx/mina Iriviaiia) for the s])cciflc 
oame, consisting of a single word; mid in Ibe Species I'lan- 
Inriim, in 1753, he carried tliis idea into full effect in Botany. 
Tlie step was a simple one, but most imjwrtant ; and Linnieus 
himself, who generally did not underrat« his services to science, 
seems liardlj- to have apprecioled its practical value.' 

704. Tbe Binomial Nomenclature in Natural History, thus 
established, Urst separated tbe name of a plant or of an aniiuol 
tmm its tliagnosiii, descriptive phrase, or character, and reduced 
the appellation to two words, the first that of its genus, the sec- 
ond Uiat of its siieuies. The generic name very nearlj- answers lo 
lite sumanie of a person, as Browm or JuntM ; the ^Kciflc answera 
lo tlie baptismal name, as John or JainM. Thus, Quereut tdba is 
tlie botanical appellation of the White Oak ; Qiierciu being tli&t . 
uf tlie genns, and ailia (white) that of a particular species : wltUe 
tlie Red Oak is named Querent rubra ; the Scarlet Oak, Qutrcut 
eoceinva; the Live Oak, Querrui virgin : the QnrOak, Querrat 
macroairpa: Magnolia ijrandijliira is Lai'gC'flowereil Magnolia; 
M. macrop/iylla, Long-leaveil Magnolia, and so on. The name of 
the genus is asnbstantive. or atleast 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.* This com- 
bination of generic and spccillc name is the name of tlic plant.* 

70j. By this sj-stcm, not only is the name of the plant reduced 
tfi two words, but a comparalit'cly moderate number of words 
serves for the complete designalion of more than 120.000 plants,* 

' MoreoTCT, he may be mid lo Imvc adoptpd ralhcr thmi originalwl iho 
idea; fur lingle-wonk-d siHicific iiaiiivs were lued lialf a eentur/ prciiona 
bf Bauliiuann. o/i'ui Rivium. 

* It la Id be nolo! tlint liic eUtsicBl Lklin names of tmi are all fpininint, 
tlicrtforc Qiieifiu alba, P'mia rigida, if. 

* Tlic name of a tubgrnui is Homciinm written in Wtwpen ilie ttro parta 
of llio plnni'a name, ai Pninut { PtuUt) Vi'glniana. Thli ii Ilic name of Ih* 
plant aiid Mimetliiiii; mori>. In aildiiioii in tlic name of Ihc ipH'In. tbat of 
the variety or ven lubvariirly ji (omelimi-* addod. 

* AlplionH.' DeCandnlle aevoni] yean aKiiex'imaitil tlx' kniwn •porfra of 
Flowering riantt al Uftwivn 100,000 anil 130,000. Tlic Inrgor numlxr imr 
pcrhap* ini'lurle llie higher order* of the Flowerleti tprii'S. In llie pnncr.t 
■taie o( our knowlHK'' of th<^ lower nnlm nf Cryptoganu, no close ndmale 
can b« wt^ll fanned of llie aeloal nuniber of ipccjel. 


in a manner which avoids conflision 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 tlie same genus, but may l)e useil in any other genus. 
Tliey are so widely thus employed that the number of si)ecific 
may not exceed that of generic ap[>eliations. 

70G. 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 indisi)ensable, 
and conformity to admitted rules is a manifest duty. Such rules 
were systematically formulateil first by Linnajus, 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 impoilant of them remain in full 
force, while man}' of the more particular rules restricting the 
choice of names have been abandonee! . The code was judi- 
ciously reviseil (in his Th6orie Elcmentaire) by DeCandoUc 
" who was ruled by the idea of having the law of priority proj)- 
erly resiwcted," was critically considered by Lindle}- in his In- 
troduction to Botany, and has of late been reformulated by 
Alphonse DeCandoUe under the sanction of a Botanical Congress 
held at Paris in 18G7.» 

707. Roles for Naming Plants. These ^^ should neither be 
arbitrary nor imix)sed by authority. They 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 

^ Lois dc la Nomenclature Botanique, etc., Geneva and Paris, 1807. In 
the Engliifih edition, translated by Weddell : Laws of Botanical Nomenclature 
adopted by the International Botanical Congr4»s held at Paris in Au^rust, 
1807. together with an Historical IntrcMluction 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 Soi.*iety of Belfdum. See like- 
wise American Journal of Science and Arts for September, 1870, and Auf^ust, 
1877 ; also, Bentham in Journal of the Linnean Society, xvii. 180-11>8, in which 
a just distinction is indicate<1 between chanfnn^^ a well-established name and 
giving? a new name to a new plant. See American Journal of Science for 
April, IH70. 

Mention should also Ix* made of Strickland's Report of a Committee on 
Nomenclature to the British Association in 1842, of Agassiz's classical preface 
on the nomenclaturt* of genera in his Nomenclator Zoologicus, and of Dall's 
thorough and welUligested Report of the Committee on Zoological Nomen- 
clature to the American Association for the Advancement of Science, 1877, 
— these dealing primarily with zoology. 



tliat may create error or ambiguity, or throw confusioo 1 
sc'icMW. Jftxt in importance is the avoiilaiice of any uselc«a 
iiitrocluetion of new name*. Other eoDside ration a, siicli as 
abaolut^i gruiumatieal c-orrectness, regularity- or eii|>liuiiy of 
names, a more or leas prevailing custom, respect for iiei-Bons, 
&c., notwithslAiuling their undeniable Importaneu, are relativity 
acuesBory." (AlpU. DeCandolk, 1, c.) 

TOM, 'i'liu following ore universal rules in scientific nomcn- 
elnluru : — 

1. Names must Ik in Latin or be Latinized. Those from the 
Greek (which an> more and more abundant, uning to the fueiUty 
oflhis Inngnugi! for coui|>oun(ling) take Latin form and Icmiina- 
tion.' Those from modern or other than classical langnages 
should at lea^t have a Latin termination.^ Hybriil Dumcs, 
nanielj-, those roruie<l by the combination of two languages (at 
least of Latin and Greek), should not \k made* 

2. For eaeh plant or group there can be only one valid name, 
and that always the most ancient, if it is tenable, 

S. Consequently, no new name should bo given to an old jilaat 
or gronp, except for necessity. That a name may be bettered 
is no valid reason fur changing it. 

709. Nmum of CeiierB are substantive and singular, of one 
woiil ; oiicl the same name cannot be used for two genera of 
plants.* They may be derive<l from any source whatever, from 

1 T1>ii«. wonUmiling wlilillie Givek « gcmvrnlly cliangclt to Ht.and wiih 
ot to Mm. A riilo tiul always oburvoil ; for while wo linvc Eiiidtudrum and 
Ori/dmitnim, Llnneiu liinuclt variously wrolc Li'nWrniJnini mid Litiadrmdnit, 
HlifHMnidniiii anil Rhadodrndrm -. and llje Greek fiiriti now pn'vaiU. 

* In tills ai In olhvr caii<i. ionic exceptlont are well iitnbliilicil bjr 
euitoni. but they oushl nol lu be extended. Tile rule as lu Uitliiiiatiua b 
miricinl u rrtpceu arlU«({nptiy liy tlic Dcveielly of prcuTvin); mudcm 
conimemoratire namo* In a recognizable form. 

■ Rut we cannot eliange nutnemu* old lumes rnr IliU fault, auch *i am- 
tnJi-ido'ilrt. f«matio'ilrt. rmuBnlouln. and icirpoiilrt (llioutfli lllpy ought W 
Imvcbecn MniW'WiM, nuniiinifinn,Bndj>rii7u'iM| ; and nnHlirn botanltl* lure 
nnt fcrDpled tn append llw cxpreiriTe and convenient Greek lemi -aldn {Of- 
nifyln; likeneiis) to irencrie names not of elaMi(«l origin Ex. aliulilaldm. 
bin!dM.JantdUmdrt.fivkiii^dri,i)tnthii<»diM,liibflltiidrt.loiinirfi<r1liHd't. In Bat- 
Ibh. lomc tiyhrida will p»nielu«te themaelTes, a* (or instaace ttnmtiiJt^, 
notimfin. millimrlr*. btatirorrors. 4e. 

• Very many, indeeil. are ailjectlvca used as snImtantiTes, as Amarh, 
Clararia, Saptmaria. rmpalient. Tricniidit. and even GI-«rigK\. AfiniUli, A«. 

Snme iwn-worded jK'nerk nmnei pnsterinr to LinniriiE, aiu'ti oi iMiti t.rtm!t, 
T7i;» Idt'ii, Buna PnHorli. remain fir leelions and «|ipeies. but not for gen- 
era. Wlien two wnnla are eonfluent into one, they are not objeetlonahln 
■■ IjiHmrnuu*, Carirmama (commetnoraling Chariea Lumau, Carolu* Ii»- 
m»nw). &«. 


prominent or peculiar character or appearance, frnm localities, 
from the names of .pereons (especiall}' of discoverers) , fi'om 
indigenous or vulgar names, or even from arbitrar}" combina- 
tions of letters. Unmeaning names, if not in principle the best, 
are never misleading. The main requisite is that the}* jshould 
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 Sauguinaria for an herb with red juice, 
H<Bmatoxylon for the Logwood tree, LUhospermum for a plant 
with stony seeds (or seeming seeds) , Myomrus for a plant with 
g3-noecium resembling the tail of a mouse. Names of this sort 
do not alwa\s hold out well ; for Chrysanthemum^ so called from 
its golden 3'ellow blossoms, now has many white-flowered species, 
Polygala is wholly destitute of milk, and many species of Cbn- 
volvnlus do not twine. Neat anagrams are not bad, such as 
Brown's Tellima for a genus nearl}' 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 every 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 accessor}' particles (thus Candolha^ not Decandollea) ^ takes 
the final -a or -ia and becomes feminine ; and its orthogi*apliy is 
preserA'cd as far as possible, making only necessar}' concessions 
to euphony and to the genius of the Latin language.^ 

The Linnaean canon forbade the use of the same generic name in botany 
and zoology, — a rule now imi)088ible to maintain. Perhaps we cannot pre- 
vent the duplication of phsenogamous names in the lower Cryptogamia. 

1 Thus, we may write Lescuria instead of LenquereuiiOj although Michauxia 
is the form for the genus deilicated to Michaux, however pronounced. The 
genus dedicated to Strangways is written Stranvasia (although Strangvcaysia 
might have been tolerable) ; to Andrzeiowsky, Andreoakin : to Leouwenhoek, 
[jevcnhookia (although the elder DeCandolle restored all the vowels), &c. 
As specimens of overdone simplification, there is Gundclia, named for Gun- 
delshoimer, and Goodenla, named for Bishop Goodenough, although Gundeis- 
kekmera 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 Lechenmiltia is />»- 
chfnaultia^ Nuttall's Wisteria (named after I)r. Wistar) \% Wistaria. The 
rule laid down in the code as drawn up by Alphonse I)eCan<lolle is: 
" When a name is drawn from a modem language, it is to be maintained 
just as it was made, even in the case of the spelling having been mlsunder- 



710. llir rtyifi'flugy of a Dew gonus shoiilil alwaye be ^n 
or the LinoitaQ refttrictiuiui. unr Imlils, viz, that tlie Daint» of ' 
genera an.- uot to l-ii>1 in -oidi, as many of ibc oIiKt names <!»). 

71 1 . NaiBM of ttpMiM ai^- (fjoimDiilj and by j>ivfortin» tuUuo 
tivM, agnving with tlie name ot the genus, and espr^ssiw of I 
MuoK.' cltaracter, habit, mode or \i\atx of growth, time of flower- 
ing, ur (.■oniineinoraliiig tin- diwoveivr. tirst describer, or aoioe 
one- iiUienrUe cunn<-(-t«d with its hislor)-. Thus, in Uie geons 
JlanurtmlttS, li. butbotii* is nameil ttom tlie bulb-like crown or base 
of the §t*iii ; R. aerU, from the* acridity of the juice ; It. jm(^ 
ratut (the accursed), in reference to tlie sanie projierty ; R. rtpnu, 
Twm tin- enT-|)iiig habit of the fitetns ; It. patillut, ftwm g 
) iiaigni flea nee ; It. aquatHii, from its growing in water: S. i 
ruli$, Trom living near eleruul snow ; R. Penn^lvimititt, tnm , 
country or State whence it was lirst made known to botanUts ; { 
R. liimplandiimui, in honor of Oonpland, one of the discuvci 
and ao on. More commonly, when a discoverer or investigatof ' 
of a H{>eclea is comniemorutf d in Ihe name, this is a subataiiUve, 
in Die genitive, as Rannnnilut NullrJIu, i. e. tlie Ranunculus uT 
>'uttall. insU-nd of B. NnJtalliainit, the Niittalliaii Kanunculua. 
Yet tlie latter form is preferred when the s|»eeies ia named in 
honor of some one who did not <Iis(4>ver nor treat of it {which ' 
should aeldom be) ; but this distinction is a custom rather than < 
a nile, and the form of the coin me mora tive name may be settled 
by Mipliony or convenience. In any case, the personal najna 
slionld have a capital initial. 

71*2. Many s|Kcit1c nnmca arc siibstantivcs, occasionally a 
common substantive, as StflOrria nemonm (of tlie groves). Oon- 
volfulut tfpitim (of the lierlgcs). Cataia jtumilio (the dwarf) : 
more commonly it is a substantive proper name, anil thia usu- 
ally nil "I'l Ecnerie name reduced to that of n epe<.-ies. Ex. 
Jtanunciilui hlammula, R. Tliora, and /?. Cgmhalarin ; nUo Lirio- 

■lood I17 thp Muihor, and Juilly dcKTving to be n-iticisiMl," Bnt this 1| 
KimrwIiHi tiio atiiuiliili?, tincv il i« allnwvil that obriou* eirura In Ihe ooik 
tlriUTilon of niinii'* of Ijiiln or rireek ilerivHiion may be correcied. provltM 
tlw cIiunKv Anvt nni urTt-ct llu! initial letter or sellable, and thai no aadonl 
nantn an- l<> Ih- <llilurl>«l. 

Tlic olauH iliat fnrbiiU clianfT" to the ortlio^rrapli.v of ant-ti-nl namaa, , 
t-Tcn In makv llirm clawiutl. U a rc^rr iirnpcr me. The botanical lAtJn . 
nf Tnlirnofon, l.liiiueun, JumIou. atxl thtir oonli-mporarii-*. Iiai hj ]tnt- 1 
•cri|itiiin ritfliti wliii-h liolnnigli arc bound to ropn-t. Wlii-n-tor* Pgrtm 
U Ihe iMitanicnl name i>( the pcar-lnM^. nulwilhiitunr1in([ the eUfsti-al Pintt. \ 
8n litrii, ai a iprclllir nann- for a imiioih plant (and a* distinitiiiilied b 
Itvit, a Hitlii Of uliKUt unci, l> flxnl by Ions botanical lur. althonph a 
(••uit claMJc*'' --•*lll*uiini'<!(.'»*r}' tniUianginauuKi'u/uiam'ita/tiM 


dendron TuUptfera, Rhus Toxicodendron^ Dictamnus Fraxinella. 
These proper specific names take^ capital initial letter.^ Rarel}' 
such a name is in the genitive ; as Helerotheca Chrysopsidisj mean- 
ing a species of Heterotheca with the aspect of a Chr^-sopsis. 

713. Specific names should be of a single wonl. Some few 
are com|X)un(led, as purpureo-caruleum ; and some of ancient 
origin (once quasi-generic) are of two words. Ex. Panicum 
Crm gallic 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 wholl}' meaningless. A plant is named b}' 
the mention of its generic appellation followed by the specific. 

715. Names of Varieties. These are in all particulars like 
specific names. Many are specific names reduced to a lower 
rank. The varietal name is written after the specific, thus: 
Ranunculus Flammula^ var. reptans^ and R. aqnatUis^ var. tricho' 
phyllus. Varieties of low grade need not be named. They may 
be designated by numbera, or by the small letters of the Greek 
alphabet, «, ^, &c. When the varieties are marked a and j3, 
the first is supposed to be the type of the species, or both to 
be equall}' 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 onl}' vernacular names, 
at least not Latin ones such as may be confounded with tnie 
botanical names. 

716. Names of Hybrids are difficult to settle upon any com- 
plete system. When of unknown or uncertain parentage, the}' 
have been named in the manner of species, but distinguished by 
the sign X prefixeil. Ex. X Snlix capreola. Hybrids of known 
parentage are named by combining the names of the two pa- 
rents, thus : *S^. purpureo X dftphnoides, or X *S'. purpureo-daph- 
noidex^ 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 'S. daphnoideo-purpurea. 

^ In ro8p<>ct to the initial of greographical specific names, being adjec- 
tives, such as Americana^ Canadenah^ Virtfiniann^ EnroiMta^ Anrjlica^ usage 
governs, and tliis is divided. But the elder DcCandolIe, who ruled in ail 
such matters in the preceding generation, always employed the capital in- 
itial, and two generations of DeCandolle follow the example. Most Knglish 
authors until recently and some continental ones adopt this usage; and it 
acconls 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 



717. The Fisation and Precision of Names. The naiDO of a 
plant is fixofl lij' publioation, auil takes its ilate IVoin tlie time 
wlieii it ii thus made liiton'ii to liotauists. 

718. A genus or oilier groujj is publislied when its name 
and clmi-aclurs (or the differences between it and all other sneli 
groiiI>s) are printed in some book, journal, or otlier 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 eliaracters, and charac- 
ters nitbout name, do not amount to publication.' 

711>, A species is not named unless il has assigned to it both 
a generic and a specific name. It is not published until it i 
mode known, by name and characters (or bj- name along witii 
sufficient information as to its characteristics), in the manner j 
aforesaid. (718.) Adequate distrilnition, among botanists and I 
pnblic herbaria, by sale or otherwise, of a collector's or chstrib- I 
utor's specimens, accompanied by printed or autc^raph tickets, 
bearing the dat« of the sale or distribution (timt is, [lublication ] 
by named E^aieeatie in place of printed descriptions), is held to | 
be tantamount to publication.^ 

720. Characters, references to date and place of publication, 
and the like, belong to bibliography or particular ph)i<^niphy, 
not to nomenclature ; but pni|>er idontilleation of names requires ' 
tliat the name of the author and the time and medium of pub- 
lication should lie taken into account. Anterior to the binomial 
nomenclature, the iKitanical name of the common tall Ilnttercii|> 
was " Ranuneulas pralcrttit ertcliu aeris" according to BauhiD, 
in his Pinas, p. 17'J. Under the new nomenclature, which re- ■ 
duoed the spcciflc part of the plant's lumc to one word, this | 
became Rannnculm aerit in Linnieus, Species Planlanim (e«l. I ), 
p. 5J4 : and a brief character gave its distinctions. In later | 
works it has been more fully dcscribeil. in some i11iiBtrat«d by I 
figures. The citation of these works arranged in ehronologiuil I 
ortlcr (or in some order), with reference to volume, page. a»d I 
in some eases figures, is the bibliography of the plant.' A bob- | 

' Mnmes may be comtnun it's led. in mnnuBcrigit ur otlirrwioif, t)y Iho fita- I 
pounder in an Builmr nhu may muke tlinn kticivrn bjr puliUeation i tin 
dal« uf Lhe Ki'niin or otlisr group U ihnl of actual pnbliL'nllon. 

' Thia itwa nut vovcr all rho condjlioiui iit publicntton, ilneo It doe 
tpcvify the L-JiaratrierB (and the mtiio may be Mid n( a piilili«be)I H| 
Willi annlf «i<«| ; linl. an tlie other liand, it ('oiivej-i lo tlic compelcat pi 
recelviug iliv lamc all tliii information and moro: so tlial It ilioukl i 
tlie rigliu nt true publication ti agalnit any anthiir to nhoin anch n 
are or atuiuld l>o known. Tlial ii. «ueli urn not in ttiv f«tiK»ry o( " m 
liaheil nanies," wlilth Krneratly uuglit to U left nntoucliid, 

* For good ejUBivlci of biblioEtraphy, arc luch detailed works ■■ 


anist, in referring to this or an}' other plant, might cite any work 
wliieh describes it, or none at all. Bununcuius acrts by itself, 
as it hapi^ens, would lead to no ambiguit}'. Not so with many 
names. For the accurate indication of the species, it is generally 
needful, or highly convenient, to specif}' at least the name of 
the author who first published the adopted appellation. 80 we 
write Ranunculut ucrts^ Zt/zn., or Z., the abbreviated name of 
Linnaeus.^ Here we have the name of the plant, and the bibli- 
ograph}' reduced to its initial. To this, further citation and 
other references msiy 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 clearl}-. 

721. When a species is said to be of Linnaeus or DeCandoUe 
or Bentham, it is simpl}* 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 may have 
named it differentl}-, and even earlier. The earlier name may 
have been discaixled 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 fVom 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 sixscies 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 b}' Linnaeus Ascleptas Syriaca, As this plant is not 
indigenous to any part of the Old World, and does not at all 
inhabit S^ria, this trivial name is not merely faulty but false ; 
so it was changed b}' Dccaisne intoM. Comuti^ in commemora- 
tion of an ante-Linna?an botanist who collected it in Canada and 
gave the first account and figure of it. As an instance of the 
second, take the prett}- little vernal plant Anemone thalictroides^ 
Z., 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 firet 
done in Michaux's Flora ; and so the accepte<l name is Thalictrum 

Caiidollc's SyBtema Vegctabilium, and Screno Watson's Bibliograpljioal 
Index to North American Botany, in the Smithsonian Miscellaneous Col- 

1 For Abbreyiations of Authors' Names, see 385. 



anemonoidet, Mtehx.. meaning an Anetnono-likc Thalictram, and 
Hittinus is the aiitliority fur Uiis name. The names wliiob for 
any reason are stinereedeil lieixiine Si/nimgmf.' (7i3.) 

72'i. A later author may eireiimscnbe a species or a geoos . 

(liflerently from the originator of the name. To a greater or I 

iees extent, this must continually happen in the course of time. I 

Bnt " Rieiniu r-imtuunit. I.iiin.," stands nnmovcd by tho sub- I 

sequent admission of various species {known or unknown to I 

Liiiiia-tis) and tlie final reduction of ull to one liy ■ tlioruugh I 

monographer. So does Silmf. GaUica, Linn., although S. qmjt- I 

guevulnmi, Linn., of the same date, is rcdueed to it. There is I 

no sulBeient reason for writing Myosolis. liroum. or C^noglottitm, I 

Jhmcn, because tliis author restricted the limits of these gcnern ; 1 

nor to write Gilia, Henth., because Benthnm vastly extended J 

' Tlip •)-nonjmy ii nn pssdiiml part of (he blblioKmpliy or K-icnlillc I 
hUtory of n genus or epecies. Bin synonymous nnil ndiniiird tiBDie* ought I 
lo bo kepi diatincl. Keeping tliis principle in view, — ■l«o llie ileciii*«ly I 
sfflrniecl iIiK'triiie of the founder of our nomenclature, that the apecilic name ■ 
ia n nullity apart fmm the generic (to that only tlie comtii nation. of lltr Iwo I 
makes llic nan>e of tho pUnt, ai truly at the cnnililucnl hajvei make the I 
tdM'it*), and bearing in niinil Ihe [iindamenlnl importance aiul alwoliit^ I 
neu of llio rule that no new names ought to l>o made where ilicn at* Untr m 
blc old one*, — the iiutlent need not U- misled by tlie confuaittg (Iivwcnr ■ 
tpeeiiiiis) Innuvalion countenanced by m*ny xoologisti siid some baUinlat% I 
and wliieh hna ot late yean been rery fully discussed. I 

The true rule Is : " For the indication of tlie name or names of any grdnp I 
(o he accurate and complete, It Is neccsnary lo quote the author who fttvt I 
publislied the name or L'ombliuiliiin of names in queation." (A. 1M7 ) TtuM^ I 
LniUia ihalitiroidn. /jnn,. falflls llie ciindiijon, except where a reference to I 
the work aa well as Hie twme of tlie originator ot the name is dumanded. I 
Tlwn the cilation would continue. " Sprt. Pi .112," (ind might In- further «»'■ 
tended. In the Flora n( Mk-hniix, l)ii« plant was treated ai distinct hOM I 
Leoniice in Rcnns: and some ixilanists adopted tlii* view, while nthmi oCl 
equal authority did ttot. Tliose who adopt MSchaux's ^nus name tb* plant^ 
CaiJoiJiiilIiim thiliftroidtM, Uirhi. I 

Now tome naturalists quote for the spceips the author who nriglnUMi ■ 
the trivial appellation even when tram-fcpred to nnotlicr getms. Tlwy I 
would adopt the genus Caiilophyllum, yet write; C-inln/J^^laBi ikiliUmlitn, ■ 
/Jnn. Or else Ihey would avoid direct talslflealion of the fftcis liy adding I 
(«p.l, this being explained to mean thai the specific part of llie name tmljr I 
was given hy Llmueus. Tlien, a* this omits all mention nf the origiml gnf fl 
eric part of ilic name, other* add this in a parenlhexia, and write: " Oudt^m 
jA/fllaa ihiliHtnidn (f-ioi. sub Li^tier-i Midir." or " foi^a/.V'"'" IVMrJi/M 
llmlittrvJfM. fJnn. suli l^aalet," or •■ Ca«lopliglhim (Lnnticr, Unu.) tMiartUtttm 
Mifkr." All such emleavor* to mix synonymy with nomrnclalttre IVP«M» 
to lie faulty in prin.lple and anwicldy In praeiii*. In Ihc mnsl abbntvtatad'fl 
form, they slate that which I* nol true : in the other*, they impair Ihe *b»-S 
plicily and brcTliy of ihr binomial nomenclature. It is all but c*rt*ta UlM,fl 
ir Ihe iteniia faulopliylluni liad been puhUslied In tlic liffltiine of UnOBimfl 
ho wuulil iKit huvv adopted It. M 


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 the 
author who made the change should be quoted.^ He alone is 
responsible for it. But this iiile has only recentlj* been strictly 

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 contraiy. It 
must necessarily be changed when there is already in that genus 
a species of the same name ; and then synon3'mous names of 
the transferred species have their claim in order of date. But 
wliatever name is first employed under the accepted genus, being 
unobjectionable, should hold, even against an older unobjection- 
able one coming from a wrong genus. This is an application 
of the stringent rule that no needless names should be created.* 

' Thus, Potentilla Canadensis^ £., yar. simplex^ Torr. ^ Gray, and not of 
Michaux, for it is the species P. simfdtx, Sfichx. Geuniy subgcn StylipuSf 
Torr. ^' 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 I>eCandolle, 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. 
Siier(jiJarta began with Persoon as a subgenus in the year 1805, and this 
date has l)een assigned to the genus, although it was taken up as such only 
in 1810 by Presl and in 1824 by Bartling. 

^ Thus, in the case of an older specific name being known, as that of 
Chtlupsis saliijna, Don, recognized as Biynonia linearis. 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 tliinl name, Chiiopsis linearis, DC. " So, again, an Indian Grass was 
first named and described by Willdenow as Coix anindinarea, then named 
by Roxburgh as Coix barbnin, and entered in Sprengel's Systema with 
Willdcnow's character as Coix Kizniyii. All these names were defective 
as referring to a wrong genus. Brown corrected the error by creating the 
new genus Chionarhne, and selected Roxburgh's specific name as the one 
most generally known and the least liable to misinterpretation , and Brown's 
Chonachne barftata is therefore the first correct name ; for which Thwaites 
afterwards substituted Chionachne Kaniyii, 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. 107, 198, November, 1878). The following 



725. KamM of Sobgenera or of other sections of geaen bivI 
like those of gotiorik ; indeetl very many of tlicm, and lliv most 1 
fitting, arc old gtiierit: namus whiijli liuve been coiuprcJR'nded I 
in the genus by i-eduction. Unlike genera and highi-r groujjs, 
however, sections, when of Greek derivation, m&y iiiojierly tako 1 
tile termination in -oidet,^ and the typical seetiun may Iwitr tho J 
name of the genus with the prefix Au.' Seelioiis need not be J 
named at all, an^l only those of comparatively high rank should ] 

is H fiirilier cxtrat'l from Ihe same protmt againit the pnicllcc " of creatins I 
ti new name in onler to combine an old tpociflL- with a new gener 
" In h'vTus, llic wanlon multipliratiun ut ill-UefiiiMl or undc-ttnalile Kenera, ' 
Rcccinling 10 tlic varied laucics u[ (peclal bolanlslE, lias liad tlie eOcvt of 1 
placing tlK iitnie *peeU-B Eueeesslvi-ly in teveral, someiitiK* x^vcn or tJght, 1 
different genera ; and It 1* propnsed lo mainUiin for llie ipei'ifiv appsllatkni 1 
th« riglit of priority, nol only in tlie gcniu alnne in whiuh ii it plaucd, b 
in the Hhule uf the genera to wliicli, riglitly or wrongly, it liai been rvfcmd. | 
This liai iH-i-n earriwl lo sucL an exient ai lo give lo (be sjiecific b 
gencrni lUbitantive aspcei, nt if tlic gcnerii.' one* were mere adjuneta, -^ f 
a lerlou* enoroaeliTUcnl on liie bcautifnl liniplieiiy of the l.inninan nomm 
clnture; and it it to Ixf feared thai tliere Is a tendency ia ttiat direciion ii 
phsnogiLmiu botany. When a iHitaiiial diamembers an old genui, tlie rule I 
requires that be liiould itricdy prracrve llie old gpiM/iflL- names in hii n 
genera ; and, wlien ho hne wnnlonly and knowingly neglected this rule. It I 
may be rlglii to correct him. Ilui where a Ixiianist bag Mtablidied wlurt I 
he believes to lie n new ipwivi, and has tlitrrfure givpn ii ■ new ni 
changing nf thia name after it bas gni into gEnnal circulation, bccatUD M I 
ha* been diu'ovcred that some otlicrr botaoiat hsd previously publiihvd It I 
In a wrong genua, ia only adding a synonym without any advantage whal- I 
ever, and is not even rcaloring an old name ; fur the specific adjpcliv* l| J 
not of Itself ihr name of a plant. ... A generic nami? is sufflcwnllj in* | 
dicfllcd liy one subslanilve; for no two grneni In the vegetable 1' 
ore allowiil to have the Hme name; but tor » species the i-(iml>ln»Hoii <tf'| 
aubsTnntivv and adjeelive is absolulely necessarr, the two-worded m_ 
name is one and indivisible ; and combining tlie lubataDlive ot one with tlw I 
Mljculivc of another la not prnerving dibcr of tliem, imt creates s 
Inlciy new name, which ought not in stand unless the previous on 
vlciinis in thcmselrrs. or preoccnpied, or referroil to a nrong gniiu. It b I 
probaltly from not perceiving tlie difference Iielween making and choBglnf I 
* name that tlie practice ohjifcled to has been adopted by some of the 1! 
among recent botanists." Bentliam. i. c. 

■ A genu* eiiuld not properly have one of lis sections called by iia own I 
name with the addition of -n/ifu or -tym*. na Astmldrs or Aitmiith. tor U I 
1* senselrss to declare rhal en Aster renemhlea an Arler ; but sectional n 
of this cnn)po*liion niay l>e cxcelleol for secliuna of other genera, aa ox- I 
pressing analogy or re«emliTani.'e. Latin generic name* mod for aecllonal J 
mtes properly take the nddition of -tlta. or -inn. or -attrum. 

' The prefix f.« (Greek for mueh, rery. or true), prefin'd to a 
name of Greek oriBin, is Ihe proper dMignaiion of the typical section tt I 
tliat genus, meaning the group which should tiear the generic name 
genua were dividnl. The rule against hybrid name* sliould in atftebicat ] 
exclude Ibis prvHx from Latin names, but it baa not always done so. 

KOM£NCLATU££« 357 

have substantive names. Designations, however, arc conven- 
ient for lower sections ; and the name of a leading 8[)ecics 
may be used, in the plural ; as Aster ^ section Amelliy and sect. 
Continni, Subgenera need not agree in gender with the genus 
they belong to. When written with the name of the plant, 
the subgeneric name is parenthetically inserted between the 
generic and specific appellation. Ex. Pyrus (Afalus) coronaria. 

726. Karnes of Tribes, Orders, &c. The names of all groups 
8ui>erior to genera are adjectives plural, and witli few excep- 
tions are the names of genera lengthened by some adjective 
termination. £x. From Rosa, Rosece^ RosacetB^ Rosales; from 
Myrtus, Myrtett^ MyrtacecB^ Myrtales ; from Berberis, Berberidea ; 
from Tamarix, TamaricineeB ; fix>m Salix, Salicea^ Salicine<B. 
The substantive Plantte 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 -e(B, £x. Rosea^ Phaseolea^ AniirrhinetB^ OxalidetB, 
&c. Some subtribes take the name of the tribe with the prefix 
Eu^ as Euphaseolece for that subtribe of the ti'ibe Phaseoleae 
which comprises the representative genus Fhaseolus. Tribal 
names may take the same prefix, as Ettcasalpinea for the tribe 
of the suborder Ca^salpinea; which contains the tj'pical genus 

728. Ordinal Names arc formed in the same wa}*, but with a 
preference for certain terminations which ma}' denote their rank, 
esf)ecially that of -aceet^ — as RosacetB^ Myrtace€B^ Cucurbiiacea^ 
meaning Rosaceous, Myrtaceous, and Cucurbitaceous plants. 

729. The names of what we now call natural oi-ders, as 
sketched or adopted by Linnieus, were mostl}' descriptive, such 
as Ensatce^ Spathacece^ Coronaria^ Papilimiacece^ ConifertB^ Amen- 
tacece, UmbeHattx ; but a few took their names fVom genera, as 
OrchidetB, Liliacece, 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 LeguminosdB^ CorymhiferaB^ &c. But the greater part arc 
simply plurals of generic names, such as Asparagi^ Junci^ Lilia^ 
Mti$€E^ Orchides^ Lauri^ Convohuli^' Erica ^ Accra ^ Cadi, To a 
few was given the lengthened termination in -e<F, as Polygonece^ 
Solattea^ Eerberide€B^ Cary&phyflea ; to some, the termination in 
-ace€e^ as in Cichoracece^ CampantdaceeB^ Rttbiacea^ Ranuncu* 
lacecB^ Malvace<B^ THiacea^ Cucurbitace€e. Subsequent authors 
have necessarily changed all names which were plurals of gen- 

3.j8 pkvtogeaphv. 

era ; and tUe sti-onglj- prcvoleut tendency has been to gire ttMrl 
termination in -aceee to all such ordinal names, and to restrlvt I 
this termination to orders. LiniUey insisted upon uiuking tliis ' 
an absolute rule even for names not formed from generic appel- 
lations : but tbis will not be adopted. 

731), III tlie first plaee, several lai-ge orders wliieh have Iwen 
known IVom tiic Hret by sueli ehnractetistic names as Crucifera, 
Lfi/HmiHotw (and its sultordor /'o;iiViono(r«), Guilt/era, t/mbeSi- ■ 
/ffra", Compositte, Labiala, Vupulljtrce, and Conijeree, also P<Jmea j 
and Graminea, FiUcet, and c^en Aroidea and Ficoidea, will retain 
tlifse apjKllations ; hut no new ones of the kiuil will be made. 

7!)1. Also, names fornietl IVom genera whieh do not well take 
the temiiiiation in -wete may be allowed as orders to retain their 
natural Torm in -inea, -idea, -ariea., and the like. Ex. Tamaria-, ffalicineie, St-ri/phuiarinetB, Bcrbrrideet, Lenlibitiarita. Wb I 
may prefer for the sake of uniformity to write Saiifaeftr, Btrbtri- I 
daceta. Lmlibtduriace<E, and ScTophulariacete (as we should wtilo 1 
Violacra), but tliis form cannot be insisted on. On the otlicr J 
hand, n t«iinination in -acta has been allowed in the names uf I 
certain tribes to avoid eseessi\e iteration of vowels. Thus, r<»r J 
the trilte of which Vcrnonia is the leading genus, authors write 1 
Veriitmacea, to avoid Veritouieie, whieh ends with four vowels. I 
8plnca and Staphylea are the tjpes uf ti-ibcs, for whieh the nameSf I 
If they followed the rule, would be S/tirttfit and Staphg/tett, I 
endiii); unc in live the other in four eonseentivc vowels. So«i« I 
avoid this by writing Staphyltacea and Spiraarea:. Olliers write I 
Si'ipbt/lete^ but this is onlj' the plumi of Uie grneric name. I 

732. A few orders or other givups took their naiuea long J 
sgn IVoni superseded generie names. Kx. Curyophi/Uaefm or I 
Caryopbf/Heo!, Ontigrarrtr ov Onagran'ea, and Lfnlibularita. I 

733. NamM oT Cohorts aiv liistinguished by the termination I 
in -ales. This was pro|Mised by Lindley. and is adopted by I 
Itentham and Hooker in the Genera I'lanUmm. Kx, Rtmalt; I 
Parieln/rt. Mititdei, Jiutalea. Pafsifliirnlts, &e., most of tbetn I 
founded on the unnies of representative genera ami orders. J 
Euphony n^quircs some to take other terminations. Ex. Pkify- I 
gallntt, Varyopkj/Uinm. I 

734. Nmwh of ('laii»M and other groat divisions are plurals, I 
eltlicr adjective or adjcotivn noiina, expressive of the leadbif; 1 
charaeter. Ex. Polypttnl^, Gamoprtnla, AjM-uda ; Anyiotpermm I 
and Gymnotpenait ; DieofyUdimes and Monocoli/fedane*. TIa fl 
names of the two great series or sul)-kingiloms. fiiltowiiig Um I 
analogy of tlie Linniriiii elnssi-s, end in -to, and are P/iteti9g<mia% M 
or PhiinerwjniHiii, and CryplO'jamia. fl 


Section II. Glossology or Terminology.^ 

735. This is nomenclature as applied to organs or parts and 
their modifications. The actual botanical terminolog}' 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 pubUshed in 1813) is still classical authorit}*, and 
until recently has received few additions as regards terms need- 
ful in phsenogamous botany. 

736. The fuudamental 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, yet synonjms are unavoidable ; and no term ought 
to be used with two different meanings. The woi-d flower^ for 
instance, must not be used for a cluster of flowers, however it * 
ma}* imitate the appearance of one, nor for the corolla or other 
[X)rtions 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- 
Hum) is a notable instance. A bract, to go no farther, is a sort 
of leaf; and the imperfect stamens of a Catalpa-flower and 
I^entstemon 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 absolutel}- 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, diflerently modified in fruiting, is an akene in a 
Ranunculus, a follicle in Aquilegia. a Ixjrr}' in H3drastis and 
AcUea ; 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 dnipe ! Arbitrary rules cannot absolutely fix technical any 
more than ordinary language. 

737. Experience and judgment must determine what modifi- 
cations of organs should be regarded as a kind, and bear sub- 

1 Ahhough the forroor 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 r4piia, of the same meaning. 


Btanllvo InBtcad of merely adjective namea. 
elioiild mil be unnecessarily multiplied. 

T3K, Tlu> classical laugiingc of sciontitic bolaiij' being Latin, 
all tbe organs of plants and then- principal diversities arc design 
nated hy a Latin or Latinized name. Modern )angiiagu« have 
also tlieir own names and temis. Gn.>atly to ita advantngt', 
EngUsh bolAiiieal t«rminology lias adopted and incorpomtcd 
ti-rma IVoin the Latin and Greek, with slight changes, not obscur- 
ing the identity, thus securing all their precision, aitd rendering 
the simple botanical Latin of descriptions of easy aoiuisitioii to 
the English student. 

Til!). In a text-book like this, the principal names and terms 
applied to organs and their leailing modiflcatious, as also tliose 
nhU-h relate to their action (pliysiulogical tenus) , or to our study 
oftlicm (didactic tenns, such as phytography, phyllotaxy, glo»- 
**ology)> are defined and illustrated in course. There remain 
the more numerous and varied e/taraderistic terms, chiefly adjtMr- 
tivfn, np|ilicul>lc to more than one or to all organs, and vthich 
cumivosc the greater part of glossology. These, which DcC'andolle 
arranged sysicniatlcally with much elaboration, may best tie 
r(*uchcd by a gloaaury or dictionary, such as that at the end of tlila 
volume, which comprises the substantive terms likewise. 

740, From chanietcrislic adjective tenns are derived Uw* 
greater number of s|K'eilic names of plants ; of which, therorore* 
tlie glossary maj' elucidate the meaning. 

T^l. Capable us the existing system cannot in single 
wonls define all observed forms and grades, nor well avoid 
various ambiguities of meaning. Some defects of tlie firal klad 
are remedied liy combining with a hyphen two congruous terms 
to dcnot« un intermediate state. Kx. ovalo-lanceolatva, or orctf^ 
lanrmilale, for on outline between the two. Also a term may be 
qualified by the prefix tub, in the sense of somewhat, as in nii- 
n'tiittiliii. tubcordatia (somewiiat round or slightly heart-shaped), 
or diminutives (such as inlrffriuKu/ut). or su|»er!ative8 {inttgtr- 
Hmiii) or other strengthened forms (such as /MTiinyi/jt^u*) may 
lie employed. Among terms of more than one form of meaning 
are such as eali/cinat. which may mean, according to tlie contest, 
pertaining to the calyx, or of the ap[tearance of calyx ; ej/autma 
may mean in cyracs, or Itearing cymes, or in the inanner of a 
. ej-me ; and paUac«iii ma>' mean provided or beset with chafT. or 
rvsembling chaff in texture. OfU'U tbe form of the word ehoold 
distinguisli the sense ; aa/t<Hitlui, f^imished with leuvea./o&Mitt, 
with nbundanoe of leaves, while faiiitfrm may mean either bear- 
ing leaves, or pro|wiIy of Icnf-Uke texture or ap[>eurunce. 


742. Absence of an organ or quality- may be expressed by 
means of a prefix with privative signification, as indehi scent ^ not 
dehiscent, exannulale^ destitute of a ring, apetalous^ without 
petals. But the Greek privative u should not be prefixed to 
Latin wonls, nor the Latin tub to terms taken from the Greek. 

743. When the Latin preposition ob is prefixed to an adjective 
t3rm, it means obversely ; thus obcordcUus 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 
diffbrences by which the species and groups of plants are distin- 
guished and recorde<l, the structure exemplified, and the histor3' 
or bibliography 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 indeijendent of all relative knowl- 
edge, or takes no notice of relationship to other plants or groups. 
The latter intends to portray the si>ecies or group in its relations 
to others, and to indicate the differences 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 teimed 
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. 

74 G. Characters are specific, generic ordinal, &c. They are 
the differenti(B^ or marks which distinguish a group from any 
related group of the same rank with which it may properl}' be 
compared. According to the occasion and purpose, they may 
specif}' only the fewest particulars which will serve as a diag- 
nosis, or they may be extended to all the known constant differ- 
ences between two or more related species, genera, ordere, &c.^ 

* The former would answer to what have been termed differential char- 
acters, the latter to essential characters. Linnaeus divided (ij^eneric) characters 
into factitious, essentia/, und natural; by the former denoting any difference 
which may effectively diatinguiah between any two groups brought arti- 


What is now t«mio<l tlie s|)ocil1c character was the siKcilic name 1 
with LinniL-iis and his pitdet-essors ; what we call the §j>eoiUc 
LiiiDicus calletl the trivial name. (703.) 

747. Suboi'dinatiou of characters and the avoidance of ruin | 
reiu'litions require that as far as possible — regaii) Ixing had to I 
tlie form oC the work — the or<linal character should cuiiiain * 
only what is nce<irul to circuniscribe it, and to exhibit di-aily | 
tia roorpholog}- ; that the chaiactei's of trilies or other divisions I 
should not reassert any |K>iliou of the ordinal charaotGr. nor 
the geneiic character tliat of the superior groups ; anil so of the I 
sectiona and subdivisions of all grades donn to tlie siict-ivs, 
Eqimlly from the si>eoific character should be excluded cveiy ] 
tiling which belongs to the generic, or is eomtnon to its rela- 
ti\e8 generally, or has been already siwcifiect in tlie section of 
its subdivisions. !So, likewise, of tlie ^anetics under the spe- 
cies. This can be done oidy by so arranging the siiecics as best 
to exhibit their relationships, that is, by bringing togetlier or 
into proximitj' those of givatcst resemblance in all respoctSt 
or in the more imixntant respects. What these are, and liow a 1 
jnsl subonlination of characters is to be npprehendwl. eannot I 
be taught by lules, but must be learned by ex(K!rieni« and I 
IVom the critical study of the classical botanical works. No one j 
is competent to describe new plants without such study, and j 
wilhout a clear conception of the jiosilioH which a snpitosed 
new species should occupy in its genns. or a genus in its onlor. 

748. Charaetei's of orders, genera, nixl of all intermediate j 
groups, are drawn almost without exception from the organs of | 
n-uetif1 cation. In the description, these pai-ts are mostly takon I 
in order, beginning with the calyx nnd ending witli Uie ovary, I 
the fruit, seed, embrjo. But. as to the orders, some writers pre- I 
fer to preface these proper characters with a general sketch of I 
those deriicd from the vegetation, which, albeit of less syste- I 
matic value generally, are often very characteristic of particular I 
families. Rubiacefe. for example, are known by tlieir ojjfKwite 1 
entire and simple leaves an<I intervening stipules, along with S I 
fi-w floral elisracters : Sai-raceniai-ea-, liy tubular or ]>itcher-Uk« I 
leasee, along with a certain combination of a few other ehatvo- J 

flvUIIj' tngrlher. at Ihrj mitclit b« in aii nrliliuinl ki>^. miil as rcrjr unlEka I 

gi^nera nflen were in lii« avxiial nyili-m ; by llie geeiiiwl mpaning lliv dlMtnCi- 1 

lliinf, the (twcr the lietlrr. wlik'li will tpparrtte a m^up from iti ncamt I 

rvlativM ; liy tlic tliinl, nil rp«l m«rl£« of Jilfcrrn™. i". t. all nfTonlMl hy tlw I 

orgKiM of frui-iifii^Mliun, whioh onl)' wnv itkcn inin accnunl tur mtwn, Ac. I 

tTpon the uonctriicilun at tills natuntl charHPtor Linii«iu prilled blmtrlf, 1 

Riid Jtully. ThcH.' arc tlie ctiar&ctcn In hU Geiitra Pluilaruni. 1 


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 be common to other orders, though not in 
the same combination. Generic cliaracters always commence 
with the calyx or most external of tlie floral organs and proceed 
to the ovar}', thence to the fruit and seed, and end with subsi- 
diary (but often no less diagnostic) pailiculars furnished by the 
vegetation and mode of growth. 

749. Detailed descriptions of sjDecies, 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 fllament, 
anther, and pollen, the disk, if any, gynoecium and its parts, 
ovules ; then the fruit, seed, albumen, if any, embiyo and its 
parts. But descriptions of this sort in most works and in ordi- 
nary cases are partial and subsidiaiy, comprising only certain 
details supplementary to or in ampliflcation of the character of 
the species or genus. In condensed works, such description is 
wholh* omitted, or is reduceil 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 
preface<l by statements applying to the whole plant rather than 
to any one organ ; and these are given in the nominative, and 
agree witli the name in gender.^ 

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 
EngUsh. 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 disap|)ears. In monographs and floras, the desirable 
brevit}-, or such as the case admits, is secured by proper group- 
ing under a subordination of sections, subsections, and other 

^ Ex. " Nepeta Cataria : erecta, data, cano-pubesccns ; foliis petio- 
latis/* etc. In English, these adjectives without any substantive expressed 
will be seen to belong, as here, to " plant " or " herb " understood. 

' 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 b 


loi. PooetiMtloii. In |>roi>cr descriptions, and T 
oT gfMicra anil ut liiglier groutis, tlie nocouut of t-ai.-li orguu rtii-ms 
» Hi-purntp si-nti-ncf ; ami in Latin the ti-nns nrv in iIil- nomina- 
tive HUM-, except Hubaidiart' ijortitme, nliicli an: often thrown { 
iiiUi the iiiilallve. Kxccpting the latt«r part, the a^eitii'e t 
nrc ■rpnrnl4.*(l 1>)' coiumuu. A s|>ecine elianu:t«r is ulw'ii\-s in une 
flenl4?nco. In Lntin, iU l-Ihuscs arc inshily in tlie ulilmive ; and 
much diversity prevaib an to tlie punetuatiun.' ^nligciieric ftuil 
other »eetioniil ehoraeters aie cuminunlj' franitHl liko ibum of 

dUpcDwJ *tih : i-nntoqucDtl; firioiu partii-ukn mc adiJed to the char- i 
a^iY ttlilcli cjii iiui icrictly btl(iii|[ ro it. In Rcutliani'i great FIuib J 
livnili, iImi In Engli*)i, aiitvllU: i^liarnciEn arc mplaivd \iy « diantL'tt-rutie I 
(j'nopil* Nl lliv head at vauh |[iiiu> ; uud a Ivna d«*uription nailer tmA 
•pcL'ic* oomplcica llw acraiant. Morruvpr, Itcnthain, hi rvvenl worici, ■ncfa 
u III* rcvUlim of the Ocnai Caaiia, alio dial uf tlic Mimoieic, which hav* I 
Latin vliaratilvn, wrilc* Ihpic In the iiumiiiallve i-uc anil i>ai'li nK-mW in a 
■i-Iiaraie M'nicnuv, In llie dwcrlptive form, al«iidunliig itie liiug-UMiil aUa- 

I Linrupui nnployt^d only tho comnia in the apvciflu tliaracUr. along wiih 
a lubslillary uk- u( the oulun In a nianniT very unlikt ita ordinary iis« la 
pumilunllun. makhig It a [luint of leu lvalue llinii the L-uniino. Tliua, 

"CnaiiDntuiUM aluum foliii rhoniliniilco-triangularilius cnisb poatioe 
rDlegTl* : lummli ulilungii, raveinia orcctia." Spei'. PI t-d. 2. ■Ilfi, 

Hvrt, while the two main ravmbcrt of ihe >euti<ni.v arr *rpamt«d tij a 
coronu, a RulMldlnry portion nf the Knit mrnilnn'. relating to the uppHmou 
loavua, I* separated by a eolun, Liuiiiriu employed ilie colon hi th« u 
way In goiiurlc L-luracicn. Thii anoinaloua usage li now abandoned. But { 
Riii4( nuibon have folEowcd the Liiinmii palicrn in diatlnguithlng the prin? 
et|iAl riirnitHT* by coniniu only. *u dial thoM liwumi.- dte cmly puinta In tht I 
«I>vcilli; I'haractcr, liuwcvvr I'M ni plica li-d that may be. Thii», 

" IUhi^hl'ului ACNtB (Linn. Spcv. TTDj tnlUi pubeaccnllhut lUhglalirtoTc 
palniuio-partliit. luliii Inciso^lontatU avuti*. lummii linearlbui, taule nveto 
plnrilbro lubpuUeieMito, pixluiii^ulli lerellbu*. ealyce vubvilluMi, uupcUla I 
BiatiTanc lubereeio lomilnaili " IM'andoUe. PnHlruniiu, i. 30. 

Tlili li the piitH.-Iiialhiii tlirungluiul llie Pnidnirniu anil in moat nxU 
pofary lyatvinalii: wurlu- Its impcrfti^tinn la ihown in (he alnve-dted «| 
men. The primary membiTi of the lenteiiee, whii'h elianieterize tlie hMVMi I 
alein, twluuelu. ealyx, and carpels, arc dioinguliliiHi by the lanut grmile al ] 
punuluaiion whkh serves for tlt» giarti of the ftnl nioinlier, via. tlie ktbaa 
of die leavn, and for a still aubonlinale portimi. viz. Ihu fonn of tlte appar- 
most lobes. This want ut BiilwrdliiBliun is to be r«nte<lird by the osa tt 
•cmlenlnns U-iwireii tlio prini'ipal members, and of ilio i-umma» only fvr iha 
•Mvndary oni's,— a punelunlion now not uneommon. and wbich ts ailupled 
In th« Twmt flr« volunio of the Monngraphbe Plianeroganianim of the De- 
Candollcs, wliieh sapplemcnts the rrodnimns. The portlun of Ihatrdaina 
cnntributeil by Dr. Hasten bciier exemplifles this than dors the rest of the 
volume. Kiir ihe laitwr Miurilli-es llie adrantagv of ihe ebange by the tuer. 
Uun tit mmmas beiw(>vn each adjui'tiv* of a uimiDunus ablative phraa* (aa, 
".Smilnr lanrifi^ni : liinbU toliuruw oblonitls ve! ovalii.oblongl«, uorlaoi^ 
3-&-ncrviis, sul>lu> paltldloribiu," fti^.}. whirv ilicy are gi^iivrally deemed 


genera. Or the members may be united in one sentence, but in 
that case the principal ones are best separated by colons. 

753. Should a point inter^'ene between the specific name and 
that of the author cited ? The practice varies. But, if the name 
is Latin, the comma is supei'fluous ; for the abbreviated name of 
the author is supposed to be in the genitive, and to read thus : 
RanunculuM repens Linruei. 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 whole 
proi>cr to insert it. 

754. In citations, the classical practice is to separate the refer- 
ences from each other and from the name by jKiriods ; thus, 
*' Anemone cylindrical Gray, Ann. N. Y. Lye. 3. 221. Torr. & 
Gray, Fl. 1. 113,"* &c. It is becoming equally customarj' to 
separate the several citations bj^ 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 otiier autlior 
has to refer to (with more or less fulness, according to the form 
of his work) is to be sought parti}* under the admitted name, 
and parti}' in the 

755. Sjnonjmj. Tliis includes all other than the admitted 
names. Ex. Hydrapdtis of Michaux is a S3-non>'m of Brasenia of 
Schreber, the latter being the earlier published name. Nectris 
of Schreber is a synonym of Cnbomha of Aublet, the latter hav- 
ing priority. - ThcJictrum anemonoides of Michaux has for s^'no- 
n3'ms Antmone (haiictmides of Linnaeus and of many subsequent 
authors who followed him in referring this ambiguous plant to 
Anemone (721) ; and also Syndesmon thaUctroides of Hoffmannsogg 
and Anemone/la thaUctroides of Spach, who projwsed to consider 

taperfluoufl. The preferable punctuation of the character above-quoted 
from the Prodromus wouhl be 

Ranunculus acris (Linn.) : foliis pubescent ibus subglabrisve palmato- 
partitis, lobis inci80-<lentati8 acutis, sumniis linearibus ; caule erecto phiri- 
floro subpubescente ; pedunculis teretibus ; calyce subviUoso ; carpcUis 
mucrone suberecto tcrminatis. 

The advantages of this style of punctuation wiU 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. Kightly to expr(>ss Uiesubonlination of charncters, the plan 
adopteil in the Synoptical Flora of North America is rcconimendetl ; that is, 
with colons separating the principal members, semicolons for subonlinatc 
and dependent ones, and commas between the adjectives of the same noun. 

* 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. 



it an iiit(!mi«ilist« gcnns bptwecit ThAlictram and Aim 
sjrstPiiiatic works. Ihe !)»<.ific cfaanictpr immediatirly tulluwfi tit*"] 
mtni-. BU(I gim-ralli funii^ a \<&H ur tint saatt SHrnUncr ; aiwl u 
Mkiwvd flnil b}* citaliutia of aulttuis wlio trnvr ttdujitirtl tW nniar, 
antl tiicn by the sj-nonj-my, or bs miich ot it as the plan of tlw! I 
work (-alls for. Tbe synoDvmous names aoil tbe n-fcivnocs \ 
nrnlfr Ibrm bIkhiU be cited iii llie ordvr ot Ihvir iHiblii-ukn 
Ilqt, to rtunomizo space, alt the authorities Tor the same nam 
ate Ivrtmght tof;rtber into one itenteDce, ami airangml aoconling I 
to tiK-ir <late. Also, wliere the synonymy 13 not etaburabily | 
(li«l)bjie(1. the various synonyms or the same generic name ar« J 
iMunlly (itaivtt in coiiaecative onlcr. 

TAG. Inmogrvfkj. Tbc Icatling and most essential citation is I 
that or ilie autUor t)y whom and tlie wortt in wlit<-b a plant la \ 
namctl and ilescribetl. ami also llie worh in whieli it is best diar- 
act4-rize(1. Among tltc cliorncterizations, puUishtxl itgtires liohl | 
a prominent place. Tlic citation of these is an im|>ortant |HUt 
or Ilie ai-min,\7ny. Tlie best iHitaniL-al iilates an- those wliieli j 
give dcHtiltti nnalj'ses of llie parts of the dower, IVuit, and s«vd. 
displaying tlieir stnictnro. 

"iJl. Habllxt and Station ore recorded in a sentence or pai«- I 
grnpli f(>ni>wiiig the nume, character, and synonymy of a 
The liob!lali"H In the place, 'lislrict. or region at ur witliin whieli \ 
till- plant iw known to lie indigenous, or to grow sgiontaneously-. 
I'he complete linliitat is the get^cniphieal range. The tiatian i» | 
tlip situation it affects, wlietlicr in water, in marshes, on shores, I 
on hill" or mnnritnins. in furesls. on ojwn plains. &e. 

I'lH. IHHTOTcror, Ac. To Ihe habitat and station of newly I 
(tiseovereil, rare, or local |>lnnle siionld be ap|>end(.fl the t 
of the diseovcrer or the eiillectors by whom the s|K>eie« lias be- | 
come known to seienee. ut least when the plant is tli'st pnblislietl. 
Date of discovery shonld also tlion Iw in'lirated. 

7.^9. Time of BtossoBiltiir should lie iiTonliMl. either the moDlh 4 
or the sc.ison, to whidi may lie added thot of the maturity of tllc I 
fruit. When the month or season is mentioned wiiltoui fartlter I 
rxplnnnlion. Ronei-ing-time is intende<l. In a llm-n. this may I 
sometimes Ix- iniliealwl nnder the gemia for all the species. J 
In the flora of an extensive region, and in rc8|»oot to B]MXiic8 ai ■ 
eonsldenibk' range in latitude o|- longitude, tlic time of fl 
(liOers so widely at the extremes of the geographical range that] 
it cannot well be si^ecilicl exe<'|it in general tenus. as tprinff T 

. &c. 

760. Etfrnolo^jr of Xamei. When a new generic name ia pnb- 1 
lished. its origbi ami meaning should always be given, If tlw | 


nature of the publication will allow it. So likewise of species, 
e:;cept where the source or signification of the name is mani- 
fest. This is commonl}' the case as respects most characteristic 
specific names, and also those cb-awn from station, habitat, and like. 

7G1. Aceentnation of Names. The pronunciation of botanical 
names is settled by the nilos 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 
falb. 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 iienult is a long syllable, it takes the accent ; when short, this 
rewdes to the antepenult. The accentuation may accordingly 
be suinciontly indicated by marking the quantity of the i)enult, 
either lonj^ as in Erica, or short as in Arbutus and Gladiolus. Or 
else the accent ma}* be marked b}' a projjer sign, as En'ca^ A rbulus, 
Glad lo! as. An endeavor has been made to represent the longer 
soiuul of the vowel by the grave accent-mark, as Erica^ and the 
short by the acute, as Gladiolus. But this plan is encuml)ered 
with practical difllculties. 

7()2. Abbreriations 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 tlie customar}* abbreviations 
in the citation of volume, page, plate, &c., in which there is 
nothing peculiar to botiiny. 

7C3. 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, fur Linnanis, Juss. for Jussieu, Rich, for Richanl). 
Sometimes moi*e of the name must be given, in onler to distin- 
guish those beginning with the same syllable. So we write Mlvhx. 
to prevent confusion of the name Michaux with that of Micheli, 
which, being the earlier, claims the abbreviation Mch.^ and 
Jkrtol. to distinguish Bertoloni from Bertero. vSometimes a 
• much-used name of one syllable is abbreviated, as Dr. (or 7?. 
Br.) for Rol>ert Brown. Initials or abbreviations of the bap- 
tismal name are needed to distinguish l>otanists of the same 
name ; as P. Browne in distinction from Rolx?rt Brown, Ach. 
Rich.^ Adr. Jiiss.^ Alph. DC, to distinguish the younger from the 
older Richard, Jussieu, and DeCandolle. Or. where father and 
son, the abbreviation for the latter ma}- lie Juss. f I., Hook. fil.. 
or Hook.f.^ &c. Certain, but veiy few, well-known and eminent 


Dames are al>brerialed lo a sign : oa L. for Unnima, DC. tatM 
I>(<'an(tulle. HBK. f^>r ilumboMt, ltoi>!>la»il. and Konth. Uwl 
bllor too long atlcr ordinarj- ahbrevialiou.' Care slioultl bul 
InkL-ii III aflis llie jjeriod by ntiich abbreviations may be di*- ' 
tiugiiuhed frum TuU naim-s. sut'b as Don. Ker. Jifytt. 

7G4. Abtirevialions of titles of frorks fuUov thi? some roles as 
those of names, or at least arc in no wbe iiec-uliar in botany. 

765. Abbreviations of the names of organs follow the aoii 
rule: Cat. for calyx. Cor. for eorulla. Siain. for stamen 
stamina, /Vtf. for pistillum or pistil, fr. for fhiL-tas or fViiit> 1 
Per. for pericardium or jjerioirp. Sem. for semen or seed, are tbe J 
most common- Nab. for habitat or geographical station. Herh. I 
for herbarium. Gen. for gi^nus. i^. or Spec, for siiecies, Var. far J 
variety, and Uie like, every one will understand. But i 
abbret iations which are common in l>olauicaI writings, at least J 
.tltoee in Latin, may ne«)l explanation to tlie eloroentarj' sliKlent. f 
A list of abbreviations is npjiended. See p. S90. 

76G. Signs. Under Ibis head might be ranked such abbreria- 4 
tions OS V. !■. for nWi tt'vam. r. ». for viJiticcam. to note that ths I 
writer has seen tlie plant, either alive or in a dried s|>eeimen : or, T 
more pailiwilaily, c t. *., when it is a spotilaneous specimea] 
that has been examined iu a. dried state, and c *. c., when it waa | 
a cultivated Kix-cimcn ; r. v. c, when the living plant was a 
in a garden only, and v. v. ».. when the sponlaneonsly gruwingl 
plant was seen ali^e. There arc also proper signs, of which tlis f 
most common are those which indicate the sexes of blossoma, f 
the duration of a plant, and the like. Also the intetTogatioa 1 
point (?) UBe<1 to express doubt; tlio exclamation ]>uinl ( ! ) tol 
indieatc the certainty that is given by the actual sight of aa 1 
authcniie original six-cimcn. Seep. 391. 

767. The marks usetl to indicate the subordination of aectiom | 
anilcr a genus, or in tlie synoptifal arrangement of genera, i 
tlie like, ore not settle] by any fixcti ndo. An approved ai^J 
rnngeinent is to employ the following marks in llie given orrtcf, f 
9 •■•-** —. Till' first one. for sections of tlie highest onlw, I 
takes numerals nfler the sign. Ex. S 1, and so on. Whoa | 

' A» Alpli. IVCaniloUp rcnmrlci, the proper ahbrcTiiition of the ni 
be«n U Cnml. But the form hf. v»« v^ry psrlr ailopipd by the first <t(l 
thr illiulrioiu name-, and liat be'/n i-iintiniint for atmngt ilin?? quartets of mm 
century. Alplmtino DeOanilntlp wf.ulil prefer to write it DC. lint In 
•ilopiwl iliat mrHle, iinr "houl'l we; for DC, ami HBK. are ennrenient a 
bri' Tint Ion* ri'iluonl to i)ir>i>. But lui'li fomia ilionlJ not be ItiLTeaw 
onlinaiy nameg Iticy woulrl be iminielKitlble. 

Kaiiiea whieh arv noi ton \onf. nnil of which an abbrevlalinn by than 
nary rule U inauffl'''-*-- incb a^ Dcaiimr, thnuld rather be wrlllen In full. 

SIGNS, ETC. ■ 869 

such sections are followed by a substantive name, they are 
equivalent to subgenera.* Ex. Phacelia^ Juss., § 1.- Euphacelta, 
f . €. the true or typical Phacelia ; § 2. Cosmanlhus, Gray, &c. 
Sections next in rank to these are marked with asterisks, ♦ fcr 
the first, • • for the second, ♦ ♦ ♦ for the tliinl one of the same 
rank. Divisions of these have the -h- prefixed ; and so on in the 
same wa}'. Still farther subdivisions may l)e 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, Monogrraphs, &e. A systematic work describing 
in proper onler the plants of a countrj- or district is generally 
called a Flora. A Flora of a small district takes the diminutive 
name of a Flarula. A universal work of the kind when it ex- 
tends to the si)ecics is a St/stem, Systema VegetcJnUum or Systema 
Regni Vegetahilis, The latest completeil Systema Vegetabilium 
is that of Sprengel (1825-1828), in five octavo volumes, on 
a ver}' condensed plan. A comi^endious Flora or Systema is 
often tenned a l^^rodromus, literally meaning a forerunner or 
preHminar}' work. But, as even this is more than most l)ot- 
anists are able to complete, the name of Prodromiis is now 
api)lied to works which are not intende<l to precede fuller ones 
b}- the same author. The principal work of this kind is the Pro- 
dromug Sytt. Nat. Regni Vegetahilis^ commenced by DeCandoUe 
in the year 1824, continued b3' his son Alphonse DeCandollc 
(aided by various Iwtanists) to its close in 1873, down to vol. 
xvii., or essentiall}' twent3* ver}' compact octavo volumes, these 
earning the work only through the great class of I)icotyle<lones. 
But the publication of the monocotyledonous onlers has com- 
menced in a series of Monographs (Monographi<B Phanerognm- 
anim). A Monograph is a systematic account of all the siwcics 
of a genus, order, or other detached group. 

769. S|)ecimens 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 P^nglish, those of the 
latter authors, and in the United States the better-known 
writings of the present author, esi^cially the later ones, may 
be referred to. 

1 DeCaniloUc in the l*rodromu8 employ e<l the wonl Stct. iSedio) for 
what answers to sub^*nu8 or at least to the highest grade of sections ; then 
§ 1, § 2, &c., for the next grade below subgenus ; and tlien the asterisk, and 
other marks. 



Sectios IV. SpECistEsa ; Dibections for their Exam 


770. ImpleqientH. Those nccesaarj' for the GxamiDatiut) of 
pbn^nogamoiis [ilanU, Ferns, and the like, arc a simple pocket { 
liMie, a siiDjile disaccting microscope ; also a shar)) tliiii-bbuleil 
knife and some needles of various fineness, mounted in han- 
dles, for dissection. 

771. Fora sing:le hand lens, one magnifji-ing only from four to 
6ix diameters is the most nseflil. A doublet, or a jiaralHiUc lens 
of Tolles, of about an inch focus, is better, but mucli more exi>cn- 
sive. The simple atage-microacope for dissectiou need have only 
two lenses (doubbts or otherwise) with lai^e field aiKl good 
definition, one of an inch and the other of about half iucb foval 
distance; uid 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 ic the study of vegetable anatomy 
and of all lower crjptogamic laotauy. 

71i. For making thin slices, a mzor is the best knife ; for ilis- 
scetion on the stage of tlie simple mici-oscope, beside needles, 
smati sea1|)els or some of the cutting instruments used by octi- 
lists are very convenient. But an ex|*eit hand is able to do 
almost every thing with a common knife or scaliiel and a pair 
of mounted neerlles. Slender forceps are almost indispensable : 
those made for tlic use of dentists are the )>est. 

773. jLnalfsls. In the examination of an unknown plant vrilli 
a view to its detei-mination, its whole structure should be made 
out, so far as the rcaterials allow, before a step is taken to i 
ascertain its name and place in the system. In respect to Ulo | 
stem, its duration and eoneistence am) iU intuiiial Htruelure, 
wliether exogenous or endogenous, arc to l>e noted. As to thu 
foliage, Uie veiuilion and tlie pliyllotasy, also the presence or 
absencte of stipules, are most imjMtvtant. The anthotaxy or 
InQorescence is to bo examined and referreil to its ]iii)|K-r tyiw. 
In the flower, the numerical plan and syrometrj', its gronnd-plan 
and the nature of the deviations from the general or tlie family 
type, are to be considered ; also the a'stivation or nrraugement 
of the parts in the Iniil, the character and extent of coaleavenee 
and adnation ; the manner in which the anther is Itornc upon the 
filament, and its place unit mmle of dehiscence, &c. Note alao 
whether, when the blossom is hermaphrodite, the autliers and the 
stigmas mature at the same or at different iicriods. The i>bcen- 
tnlion and liic ' w and i>osition of the ovules shoiikl be 


determined. Two sections of the flower should be made : one of 
them vertical and dh-ectly through the centre, in the manner of 
Fig. 336-341, — this will display the adnation, insertion, &c., 
of all the parts ; tlie other transverse and through the middle of 
the ovary, 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 u[X)n the same specimen, and these are equally to be investi- 
gated. In fresh seeds, even those of minute size, the embiyo 
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 diy, 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 paragi*aph. Such 
diagrams can be drawn by any one with a little practic»e ; and 
they ma}' be made to express the whole floral sti*ucture, even to 
the coalescenc*e and adnation.* But in the process of determina- 
tion the student should beware of trusting wholly to his dii^rams 
and sketches without direct veriflcation. 

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 fmits may be detached and prepareil for 
examination and dissection b}' 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 unhke the living state. Consequently, 
the Herbdrium or Hartiis $iccus of the botanist is to him more 
essential than the botanical garden, important as that maj' be. 

776. Herborizing.^ The collector's outfit will essentially con- 
sist of a Vasculum or botanical box, a Portfolio^ a Trowel^ a pocket 
Len$^ and a small but stoutly covered Note-book. Some use a 
lK)rtfolio onl}', others the botanical box ; but on a long excureion 
it is well to have both. The former is preferable in most cases, 
except when specimens are collected for the immediate use of a 

^ See Eicliler's Bluthendiagramme (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 Ltman H. Hotsradt, 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. 

^H class. When well stOL-kecI with paper, it is of almo^^nUmUeH 

^^M capacity ; and most pluiits uC delicate t«sture (as many of tUo 1 

^H suialler aquatiee, and tliuse with fkigacioiia or delieatc corullus) I 

^V need to lie consigned dii-ectly to tlic pajier in wbieli tliey aiv to 1 

^V be presaed, and to be kept nieanwbile under some pressure. 1 

^M TTT. The Vateuluin ia very useful Tor holdit^ plants that ore \ 

^B to be examined IVesli, and Tor tbiek roots, lai^e fraits, &c. It I 

^H is innde of tin, and shoidd be of oval-cyliiulrieal sbajw, alwut 1 7 I 

^H ini-lies long anil 4 by 6 inehes wide, and providcil witli a light I 

^H stnip to ttirow over tbe shoulder. Tlie lid opens nearly tlie J 

^V whole length of one of ibc flat sides (15 by 4 J inehes, vilh ono i 

^H fourth inch lap), is made to fit as close as posailile, and fastcua I 

^M liy a simple spring cnteh. When no portfolio is used, a larger box I 

^M maj' lie require<l. Plants may be kept fresh in such a box for I 

^1 many ibys. For a several -days exeursion, when it b desirable to I 

^1 biing home n lar^e number of fl'esli plants, a tin eliest, made some- I 

P what after the pattern of an old-fashiune^l trunk, will be foiinil I 

2 / yty eonvenient. It should be about T*-incliea Itiug, 10 iuelit.>s I 

wide, and 10 inehes high to the top of the eonvex and Itingod \ 
lid, which forms the whole top, and to which a handle ia fitted. I 
778. A good fuiTn of Portfolio is made of two pieces of Innder's 1 
boanl coveiwl with enamel cloth, and fasteuing togetiicr with two I 
long straps with buckles. Handles similar to those on a carpet- I 
bag may Iw attadierl for carrjing. The usual size of portfolio 1 
is 18 by 12 inches, but ICJ hy 11} inches may Iw U'lter, as ] 
tliere would then Ite little danger of making 8|>ccimcns of too | 
great lengtli for tUo herbarium. (784-) Or the back may be of J 
soft leather, an inch or so in width, and a light strap anil bitcklo I 
at the fKint e<lge and at each end. The imrtfollo should coataln I 
a go<Hl quantity of folded sheets of thin unsizml jiaper. similar I 
to gi-ooci-'s tea-paper, and of a size only a little smaller than th« I 
sides of the portfolio. Ver}' thin innnilla paiwr. or what is so J 
called, is eseetlcnt for this purpose, being sulHctently bibulous I 
and ratlier strong. I 

77y. The s|)eeimcns as soon aa gatlieriil should be laid neatly I 
in Uicso folded sheets (called tptcimen theeU), and kept under a I 
moderate pressure in the iwrtfulio. The shoetA with the apeo- I 
Imciis aie aUcrwards tmnsferret! to the home jiress, but tfael 
e]>ecitueiis should be left continuously' in their sheets through all 1 
the changing of driers, until cured. Indeed, the s|>ccimen8 maj J 
well r^mnin in ttic sheets af^cr drnng, until wanted for mountltig 1 
or for exchanging. For fine siMwimens, the use of this tpeeiom I 
paper is verj- imjiortant. Many plants are so extremely dclicatal 
and sensitive that they will nut bear the least handling irtUuNltl 


curling and shrivelling, unless thus enclosed : also without these 
sheets much time is lost in transferring small specimens one by 
one fi-om one driei to another in the drying process. 

780. For digging up ix)ots, bulbs, &c., a small and shari3 
ix)intcil triangular Trowel or stout knife will answer. One of 
the l>e8t "diggers" is made from a lai*ge file. Let a black- 
smitli bend the lower half of the blade to a gentle curve, so tliat 
tlie ix)int will be about an inch out of tlie true line. Grind off 
the teeth and rc-temi^er 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 manj'.^ 

781. A Note-hook should be carried uix)n 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 

782. For most plants, the best time for collecting flowering 
specimens is in the morning, soon after the dew has disappeared. 
Vesi)ertine flowers have to be secured earlier, or at nightfall. 

783. Care should be taken to have the six?cimen of the proi)er 
size, neitlier 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 l>otanists suppose that a small sprig, containing a 
flower or two with a few leaves, will answer all purix)ses 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 alwaj's the most important one. In various 
genera and orders, the fruit is the most distinguishing character- 
istic, as with the Potamogetons, the Cruciferae, the UmbelHfeiw, 
and the Cyperaceae. With many 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. Consequentl}', 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 

* [There is an EngHsli 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 vits riveted fast to the steel. It is nearly impossible to 
break it.] 


flowers and Aiiit in the same plant, or fruit may be fl'eqanQ^'l 
obtained from more atlvancwl plants at the aanie time. If not, 
fruit must be collected Inter, as in case of shrubs and trees, of 
which genemlly only a branehlet with flowers, or witli flowers 
and leaves, can be gatliered first. But suhscqnentlj- the frnlt 
and mature leav<>s, sliould alwajs l>e taken, if practi(»ble, from 
the same indiviilual as the flowers. Of dioecious shntbs or 
trees, like the Willows, each spetiics should be represented by 
four pieces ; first, the sterile and fertile catkins will liave to l>e 
oblnined, nnd the res[>ective individuals marki-d, so that later 
eorrcsiwnding twigs with mature leaves, stipules, and fruit may 
be gathered, and the specimens rightly matehed. 

TM4. A speciinen should be so arranged as to l>e no larger 
when pressed than can be neatly mounted on the heriiarinm 
pai>er. A slender plant not over three feet in height should 
generally be presened entire, root and all. This can be donu 
by bending or imitially breaking it at one. two, or three places, 
and doubling so that the sections will not rest upon each other 
• in drjing. If broken twice, it may be neatly arranged in Ihc N 
form wlieti put in portfolio. Vcr;^- large herbaceous planlft will , 
have to lie (lividcd and the parts prvsened sei)aratcly, or. IwltiT, 
take a suitable jrortion of the iip|»r stem, having leaves, flowers, 
and IVuit, and a convenient part of tlie lower stem containing { 
radical leaves and with it sufflcient root to show whellier tho i 
plant is an annual, biennial, or [x^reimtal. Thick stems, roots, 
tnl^re, bulbs, and the like, should Iw divided or thiimed down 
with a knife, but in such a manner that the original stiai>c con 
be easily mode out. 

785. Carices should be always collected when the IVuit is fhll- I 
grown, but not so ripe as to fall away. 80 also should other { 
Cyi>erace(B ; yet it is well to collect also earlier speirimcna of 
these in flower. Grasses, on the other hand, should genenlly 
l>c collwted soon aftt-r they come into blossom. For when 
matiue tlie spikelets in many sj>c(-ies hivak up and full away in 
ilryiug. The culm, leaves, and root of Stilges aud Grnasra 
should be pi-eserved. as well as the Inflorescence. The root i* I 
no less Jmi>ortiint. Cespitose sjiecies should l>c so collected a 
preaerveil as to sliow tlic tufted ciiaracter. The culms of most i 
setlges and grasses act stubbornly when bent for arrangement I 
In portfolio or press, and arc not disposed to stay in place. Thie J 
(litlleulty is jtromptly remedied by crushing with the t«elh tbo ' 
angles mndc by the bending. Or these may be Ihnist tlirough | 
sliu of pa]>er. In drjing Sedges and Grasses, very moderate I 
pressure should be employed. 


786. Somd aquatic plants (Algae espedallj) arc so soft and 
flaccid that, to secure them in their proper shape, they must be 
placed in clear water and floated out b}- inserting beneath them 
the pai)er 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 ma}' be laid directly on the siXK;imens to prevent 
their sticking. Also \iscous or glutinous plants which are liable 
to adhere to the sheets enclosing them may be sprinkled with 
L3copodium spores, powdered soapstone, or some similar sub- 

787. The name of the plant if known, but b}* all means the 
locality and date of collection, with any other descriptive re- 
marks regarded necessaiy, 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 localit}* loses much of its value and interest. 

788. Drjfng Specimens. The chief requisite for good herba- 
rium s|>ecimens is the extraction of the moisture from the green 
plant as rapidly as possible under a pressure which obviates 
brittleness. This is to be aflfected b}- pla(*ing the thin sheets 
containing the specimens between layers of bibulous pajxir, called 
driers, and appljing 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 dr}ing paper made for the puqwse. Equally good 
driers are made of the thick and felt-like brown paper which^ 
after saturation with coal-tar, is here largel}- 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 newspai)ers or of an}' 
soft wrapping pa|^r, cut or folded to the proper size, and 
stitched (ver}' expeditiouslj' by a sewing-machine), or Joined b}- 
eyelet pai)er-fa8teners at two c»omers, 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. The}' 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 lai-ge. 
For the plants dry better in small sections and with the pressure 


evenly distributed. Hence U U best to have these secdons not 
over five or six. inches in thickness, nor shouitt the jiile itself ' 
be carricfl too high, never exceeding two feet. Puinteil binder*' ' 
boards may be used, instead of tlie eommon boai'ds, to sepai'ate \ 
interior divisions. 8onic botanisU use a kind of hittice niadv of | 
two layers of thin strips or laths, crossing each otiicr. This is | 
said to allow fVcc est^ape of the moisture hy cvaiiorution, luid so 
to accelerate drying, as in tlie case of the wire press. 

790. For giving pressure, various ways havu been contrived. 
The Sereic-prtM is convenient and compact, but objectionable, J 
because it does not follow up the pressure as the plants shrink | 
in drying. This objection does not aiiply to the Levrr-pre**rt, but | 
tliey are usually unwieldy. Fortunately, onu of the best forms 
of the dr>ing-|ire38, as well as the simplest and clieapest, i« i 
merely a boartl with weights placed on the top of tlie pile of 
specimens. Here llie pressure is continuous, constantly follow- ' 
ing the shrinkage of the plants. The weight on a ]>Ue should \ 
vary from 25 to 100 pounds, according to the nature of the i 
Bj[>ecimens and the quantity in the press. On an average, CO ' 
pounds is sufficient for most plants. If much greater preseunt 

is used, there is danger of crushing the more delicate parts of the ' 
e|>ecimen, and thereby impairing its scientific value. For weights, 
bars or masses of iron may be used, boxes filled with sand, 
BtoTies, and the like. 

791. Specimens brought home in the botanical box must be 
placed in such thin specimen-sheets as are nsiti in iwrtfolio. 
In putting plants in sjieci men -sheets, whctlier in [wrtfolio or 
press, it is well to take some pains to spread out the specimens J 
neatly ; for a little care now may save much hiter labor. How> I 
ever, with most species, any carelessness in this respect can I 
be remcdieil at the lirst diange of driers. But there are some 
plants, previously referred to, so |>eculiarly sensitive tliat what- 
ever adjustment they receive must be given at the time they arc 
first placed in Iheir sheets. 

7f)i. Altlioughplantsean, if necessary, be kept fresh forseveral I 

daj~B in bos or |>ortrolio, on returning IVom a collecting trip they I 

should bo li'ansferred to the home press as early as )Hissible. In ] 

the transference, particular care should tie taken to slnughteo I 

out and remove all folds and crumpling of the leaves, petals, I 

IVonds, &c,, and to arrange the B]>ecimen as naturally as poaai- I 

hie, so as to show the proper habit. Both aides of tlie ttuwvra I 
and leaves should be exhibited. Plants that were j>ut dirwjtly 

into press should receive this special attention at the (trat I 

change of driers, which on this account should be made wiUdn J 


several hours afterwards. The stubboraness and elasticitj', so 
troublesome in specimens when first put in, will then have 
mostl}* disappeared, and the whole specimen will be found suffi- 
ciently flaccid to have ever}' part stay as arranged. If this first 
change is deferred longer than ten or twelve hours, the speci- 
mens of many species become too dry for making the alterations 
requii*ed. 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 dr}'ing. 
It is well to change these fragments of paper with the driers for the 
firat da}* or two : afterwards they may remain with no detriment. 

793. To have the specimens retain their natural color and 
general appearance, they should be dried as rapidly as possible ; 
and this result is best secured by fi"equent changing of the driers. 
These should be changed at least once a day for the first four or 
five days, and afterwaixls ever}' other day, 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 ex|)ose 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 ver}' often ; 
but more must be employed when the plants are thick and succu- 
lent. Uniform pressure may 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 al)out some of the larger flower-heads of the Composite, &c. 

796. The time required to dr}' 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 lialf the usual time, and 


the ftp(H*imon8 will \yo in much finer conrliti«m. An cT|#ntt««*'. 
colUH.'tor liaH no (UfDrulty in ajicortaininf^ wlM^thrr • plant i» 't*^. 
plftfly ciinNl or not, while to a novicv it in oA«*n a m»t:#r ' 
unc*ertninty. A tliormijrhh' dric<l plant can Im* u»iiall\ i*-** • 
itH iKHMilinr hay-like rattle wlu^n (liHtiiHietl : al<w» hy |*b« n«> * ^ 
plant ni^aiuHt the chei*k. If there is a mruaatioa of r«*#h>f^«. '.v 
plant i8 Htill moist. 

71) 7. If the thick leaveft of fleahr planta mrv ininrr^^l i * 
a few moments in hot wati*r, the fieri^Kl of (l4««irratiiiii «.:: • 
greatly haMti*ne(l ; but they iVe<|uently turn <iark a* a o «>«. 
quence of the immersion. The dning of mic-!i |>lantft. i%' 
|>articularly of the MomK*otyle<lona, may l»e ailvantatfitm«l,i 'i- 
I)e<lite<l l»y placing them lietw4*en fieveral driem ami inmiiic xl^-x. 
with hot ironfl. Small phinta may lie ver}* neatly drH-* I in •-«i 
books. Very lieautifhl 8|XK'imen8 may In> maiie liy pU«-iri;r t:>r 
plant jn a tall and narn>w ve«uM*l, and i>ourinfl: over it a »uflW v^;.t 
quantity of (*lean and dry sand. When the moiiituni' i« a>>«i«r*»«i. 
it may l)e flattencMl in a press. 

7i)H. In shining the driem of a collection, place tin* |o<iArr 
to l)e cliange<l »t the left haml on the table or OHinter. thr i»>-« 
pile in front with its length |)arallel to the |M*rM>n, — a |«*«:t:*« 
the most favorable for giving any needed at tent it >n in arran:r*&tf 
8ix»ciniens, — wliilo frt^nh driers may U* pla<*c«l at the ritfht \\xx>\. 
or lK»vond the piU* in fn»iit. Thus armnge«l. the ♦•iM-^-t* «»f *j»"> 
mens can Ih' rapidly sliiHiHl into their fn'-li driers. 

71*1*. The moist <lriers may l>e spn^ad out in the Min*hin-- t- 
dry,, or stning on n line in a wann ni^ini. cir in th«« o|M*n a.r. / 

n<»t t<M) windv. Verv moist driers mav Ih» thonMi':hI> «ln.«! 

• • • ^^ - 

within an h<»ur, if spread in the hot Mjnnhine. In in«'l« loi-c/. 
Wi'atliiT, thev must Ik» drie«l bv the fln». 

xy^K^. To nM>apitulate the ni<w»t im|K»rtant |¥>int» in {[••■■I *!•'••.• 
m(>n-inaking : I'so s|M><'inien-sh<H*ts to hol«l the plants tji«i.«- 
turlK^d during the wliole pn>cess of dning: um» pU'nt\ tif ::.-•• 
nio>t bilMilons dritTH. snn-drie<l and heatnl whi*n prarti«-a^>l«* : '\ ^ 
not make the piles t(M) large : make the tlrst shif\ <)f «Iri«'n* »:Th n 
a few lioiirs. at that time making nil ni-ed(*«l adjustnH*nt of \\vr 
tiaecid Hpr<'iinens : change the driers twice a day f«»r thr f.r»: 
day or twn. 

^ol. For rnlU'<-ting and pn»sc»r\ing specimens on a j-»«imrv. 
or ulifh niovini; fn»ni phuH' to platx*. S4mie m<¥lifl<*ati«in *^i tSv 
stati<»iiarv press i?* rc<|iiiNite. The TmreUiHtf-prtu mu'^t U* i^-rta- 
blc : arcordiiiiily the pn*ssun* is applii-<l by stnm;; lealht*r '•tra|i* 
with biicklrs. There should In? ihnt* straps, one ginlin;* \\k^ 
|>ackage around each end, and one len^hwUe. The toy and 


bottom, if of thin boanls, must be cleated, or compounded of 
wood with the grain in opposite directions, or very stout binder's 
board or trunk-board may be advantageous!}' 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 b}' 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 i>ortabilit3*, 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 drj'ing takes place mainl}' b}* evaporation 
instead of absorption, much less paper is required, and the trouble 
of changing the driers is saved. ^ 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 s|X!cimens dried in this 
wa}* are apt to be brittle. To use this 83-stem advantageous!}*, 
the botanist should have at least two such presses in o[)eralion, 
one for collecting, while the other is in use for dr}ing. 

803. Poisoning. Dried si:>ecimens arc liable to the depreda- 
tions of certain insects, especiall}* of their larvae. The principal 
l>est is a small brown l>eetle, Anobium paniceum, L. ; the perfect 
insect does considerable damage, tlie lar^^a vast!}' more. Plants 
with milk}' juice, such as Asclepiadese, Apocynaceie, and Eu- 
phorbiaccflB, tliosc 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. Ranunculacese, Umbelliferse, and Compositie are seldom 
spared ; while Labiatae mostly escai)e, probabl}* on account of the 
volatile oil which they contain. Even Ferns are liable to have the 
parts of fructification eaten awa}*. To a certain extent, the im- 
pregnation of the herbanum-cases with camphor, naphthaline, 
or strong-scented oils, may exclude the vermin. But safety is 
secured only b}* poisoning. 

804. The proi^er poison is corrosive sublimate, dissolved in 
strong (95 per cent ) alcohol. Drop into the alcohol as much cor- 


^ Prof. A. Wood seems to have been the first to call tlie 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 Rc&ssler, optician, at New Haven, Connecticut. 


roaive sublimate ns it wiU take up, then udd a trifle more of rIcoSi 
so as to koep (Lie soliitiuii just below the point of saUiratiun. 
Tlie stronger tlie solution the better, except tliat, at All! satnin- 
tion and %Tlieiv liopiuiisly used, nn eflloreat'eiic-e may sumeltmL-s l>e 
left on tUe Burf'iuie of tlie poisoned apcfimens u|)on the evapora- 
tion uf tlic nltohol. Some add to tlie solution eome carboUo 
acid, at the rate of a fluid ounce to each quart of alcohol. TIik 
solution may be applied with a soil brush (one n'itli no metal in 
its fastening), or by a dropping bottle, or even the s|K>cime»a 
niny be diptwd in the solution placed in a flat poi-celain diah. 
The brush (using a pretty largo and sofl one) is the most coit- 
yenient and efficient. The moistened a[)ecimen8 should be pla«^ 
between driers and in slinllutv piles until the alcohol evaporates. 

805. Thoroughly poison all spceimens before admilling them 
to tlie herbarium. It is well to poison all B|iocimens whatever, 
aa soon as they are made or at the close of the liolanizing sea- 
son, as well those inten<lcd for exehauges as for the collector's 
own herbarium. 

606. Keep all specimens between sheets of paper, or within 
foldcil sheets, not too crowded or overlaid, away fVom dust, and 
in a perfectly drj- place, so as to avoid mould. When attacked 
l>y mould, the corrosive-sublimate solution shoukl lie agiplied. 
A proi>erly dried specimen, duly cared for, should be as lasting 
as the paper which holds it. 

HOT. Tbti Herbarlam, called by the earlier botanists HortHt 
Sicdii, is a collection of dried s|K>cimens, named and systemat- 
ically arranged. It is indisgiensalilc to the working systematic 
liotanist, 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 berhaiinm may tw restricted to a particular (hniily of 
plants, made the object of sjiecial study. A general faerbariuni 
Bhould contain specimens representing all the natural orders and 
as many of their genera and 8|)eeies as iKwsilile, 

808. The form of tlic herliarium as to the size of its sheets is 
consideraljly variable. That of Linmvns is of the size of foolscap 
paper: this would now lie universally regarded as miicJi loo 
small. The principal British iierbaria adopt the size of IG} by 
lOJ inches, which is rather too narrow, rarely iiemiittlng two 
specimens of the same s[iecies of anj' considerable size to bo 
placed side by side on the same sheet. In the United States. 
lOi inches in length by llj in width isndople<l: that is. for the 
genus-eovers, the spccies-paiier being a ({uarlcr of an IncU nor* 


809. The specimens representing each species may either be 
laid within a doubled sheet, loosel}* (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}' time be 
exchanged for better ones. But a large herbanum on this plan 
becomes cumbrous and inconvenient for ready reference and 

811. The best plan in a lai*ge herbarium, and one much to be 
consulted, is to attach the specimens completel}*, by an}' kind 
of strong and light-colored glue, to single sheets, or rather half 
sheets. The sixjcimens 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 J inches of about 28 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 

812. In no case should more than one species be knowingly 
attached to the same sheet. But of ver}' 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 comer 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 ujK)n the sheet, indicating the hab- 
itat or the special locality, date of collection, name of collector, 
and any otlier 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 presen'c them, placing them 
in little pai)er pockets or enveloi)es and pasting these ui)on the 
sheet close to the si)ecimen to which they pertain. Sketches of 
parts dissected may be drawn uiK)n the slieet. Notes and mem- 
oranda received with the specimen or too extended to be entered 
uix)n 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 


or notes, should he atUehcd to the sheet ntnr to th^ m^^r^ 
they belong to. In view of this, |irintA'«l ami wnlim u%^u 
should be of small sizc.^ A ticket which extvu^U ftiir *'\ :« 

> All printing on an heiiMurium ticket aliuttlil br la plain tvpr . a»i u»'« 
bonicrt, uselessly occupying nioin. slioulil Iw aTuklrtl. If may U^^* • 
thought niHMiful, it should be of pUln liiu*s. It is n<»t ij«'«irml4c tn |i«r»«i ,tt 
the space on a ticket with teiMinite Uim*s and heading* tor haUtai. 4«'.« .^ 
collection, time of flowering or fruiting, name of eullrrtur. a»l ta» ^^ 
These particulars may conveniently be entered at the buctom uc ui^ U *-w 
ticket, as may be convenient, leaving the rest of tlie space frw fut ihm ■*■■ 
of the plant, the authority, and |>erbapt a synonym. 

Tickets for specinu*ns distributed among other botanisU may ««{« Lsit • 
head-line indicating the sourct>, sucrh as ** Bi coli- c v. rAaav ." ur * a t at u ' 
or, in English, " From tub IIbbbabicm or" the botanist m Isw oimma'Tm v* 
the specimen. The following may serve as an example of a •u»f4r i» ft#i 
for the Si*nding out of dried specimens, and of tlie way in whk^i tW tj kff 
may be filled out with the name of the plant, its babiut and stat*<A. 
of collector and time of collection. 


^/i^CUA^f^^A. C^^yhuXc m., 7^r. 

For the hotanint's own herbtrium. it is well to use a tdank tickrt witJ^ s 
pnnti*<l heading like the siKviuK'n aliove. but with the ** ex " oroitit^l 

When a considerable colUt'tion is nuule in any particular U»tanti-al r\x\*> 
ratidti or excuntton, and numerous or several *pi^*inM>ns of the «aiim- •(«« w^ 
arv jrathenil, to U* diiitribute<] among botant»tf in the way of % tihar^-v * 
olherwi-*!'. tht-M' are i*onimonly given out under numliers aifi •ith a ir-r-w-I 
heailing to A »|HH-iaI ticket. The foliowini; i* an approve«l fnrm ••{ toib a 
ticket, and of the miMie in which it may be filled up in writing ty m*«r.i£{ 
the name of the »|H*<-ies, the hK-alitv. 4c. 

Ali'ine Flora uk thk R«n:KY .Mucvtalns. 

C«U C C PARRY. i?7». 

No. J//, 

Colorado. Ki>w<>*« ^ C£u^ Ctia*^. 



inches is a nuisance ; and those of an inch and a quarter or an 
inch and a half in width and three or four inches in length aie 
most commodious. 

813. The sheets of all the species of the same genus, when 
not too numerous, or of a particular section of it, or any conven- 
ient number, should be consigned to one genus-cover. The best 
genus-covers are of manilla-rope paper, the " bleached manilla " 
such as that of which tags are made is the neatest article, but 
rather more expensive : they are in whole or folded sheets (pref- 
erably in quarter quires), accurately trimmed at top, bottom, 
and front edge to the size of 16 J by 11£ inches; that is, the 
folded sheet as used is a trifle longer and a quarter of an inch 
wider than the species-sheets it holds. The sheets to be firm 
enough should weigh 1^ or If ounces each, or from 45 to 52 
pounds the ream. The generic name should be written in a bold 
hand on the lower left-hand comer ; that is, on the upper face next 
the back : at or near the lower right-hand corner, the name of the 
containeil si)ecies ma}' be written either with a pencil or in ink. 

814. The genera should be arranged in the herbarium accoixl- 
ing to some systematic work, and numbered accordingly on the 

815. The herbarium must be presented in close cabinets or 
cases free from the access of dust. Tin cases, just deep and 
wide enough to receive comfortably the genus-covera, and about 
six inches high, the hinged lid being one end, may be recom- 
mended for a small collection, as the}' are dust and insect 
proof, are portable, and may readily be arranged on shelves. 
But, for any herbarium of considerable size and continued growth, 
woollen cabinets with well-fitted doors are to be preferred ; the 
interior of the cabinets being divided into pigeon-hole compail- 
ments, full}' 12 inches wide in the clear and 17 inches deep, and 
not over 6 inches or in small herbaria not over 4 inches high. 
Into such pigeon-holes, the genus-covers with their contents will 
slide readily, and may be compactly stowed away. An index 
to the genera of each order may be affixed to the interior of the 
cabinet doors, or pasted upon the upper face of thin boards, 
inserted at the beginning of each order. The name of the order, 
written or printed in bold letters, may be pasted upon the front 
edge of this boanl, or upon a flap of card-board affixed to it. 
Moreover, it is well to write the name of the order upon each 

816. Except in public collections, where fixed cases may be 
preferred, the cabinets should individually be small, only three 
or four feet high, and containing only two or four vertical rows 


of compartments. Such cabinets can be increased in numl>er as 
required, are portable, and can be disposed in any order, side by 
side or one surmounting another, as nia^* be most convenient. 
Tlie doors should be so constructed as to o^jen and siiut readily, 
but to close tightly, so as to exclude dust and insects.^ 

^ An excellent plan for Bmall and inexpensive herbarium cabinets, of a 
p3rtable cliaracter, id proposed and illustrated by Dr. Parry, in the American 
Katuralist, viii. 471. Each small case is in fact a plain wooden box, wide 
enough to hold two tiers of pigeon-hole compartments, and of any desirable 
height (three conifMirtments high in I>r. Parry's plan, but double the number 
might be better) : the entire front ccmsists of a pair of doors meeting in the 
centre, there fastened by a flush spring catch ; the doors bevelled on tin* inside, 
with a corresx>onding bevel on the case, to which tliey are attached by out- 
side hinges, so that in oiK'ning at a right angle there are no sharp comers to 
hinder the drawing out of the herbarium papers ; also allowing the cases 
to stand close sitle by side, as well as one upon another, without interfering 
with the free opi^ning of the doors. These, moreover, may swing quite 1»ack 
against the sides without in any way straining the hinges. For lifting, a 
pair of flush handles, countersunk to the level of the wood, may be attached 
to the sides. When the herbarium has to be removed to a distant place, 
these cases, having no projecting knobs or handles, will go readily into ordi- 
nary packing-boxes. 



Ach. = 


Berkh. = 

= Berkhey. 













C. Ag, 

C. A. Agardh. 



J. An. 

[ J. G. Agardh, son. 









Marschall von Bieberstein. 




Jacob Bigelow. 






Andersson of Stockholm. 
































Wm. D. Brackenridge. 











Brew. ^ 

) W. H. Brewer & Sereno 
) Watson. 

















Benj. Smith Barton. 



W Bart. 

W». P. C. Barton, nephew. 

Br., R. Br. Robert Brown. 


John Bartram. 


Patrick Browne. 


Wm. Bartram. 








Palisot de Beauvois. 








J J. Bennett. 



A, Benn. 

A. W. Bennett 








[ Cambessedes. 


M. J. Berkeley. 




Campd. - 

— Campdera. 

Eat, == 

Amos Eaton. 


DeCandoUe, usually DC. 

D. C. Eat 

. D. C. Eaton, grandson. 


























A. W. Chapman. 






































Cunningham, A. or J. 




Wm. Curtis. 



M. A. Curt. M. A. Curtis. 


















1 A. P. DeCandoUe. 




A. DC. 

Alplionse DeCandolle,8on. 


J. G»rtner. 


Casimir DeCandolle, the 


C. T. GsBrtner. 


Decuisne. [grandson. 





( jar id. 
















(ifrm . 


Ih amnr. 








IM so. 



(jingins de Lassaraz. 




















J. G. Gmclin. 



C. Gnvei 

C. C. Gmolin of Baden 


Dodona^us (Dodoens). 

*S. (imel. 

S. G. Gnielin. 



















Gn r. 







Dnhanid du Monceau. 





Gron. \ 




Oronoc. ! 



Guett. - 












Ilcujenb. Hagenbach. 

llaU. HaUer. 

11am. Hamilton. 

Hanb. Hanbury. 

Hanst. Ilanstein. 

Htuiin. Hartmann. 

llartw. Hartweg. 

llaro. Harvey. 

Hasa. Hassall. 

llassk. Hasskarl. 

llmtsin. Hausniann. 

Haw. Ha worth. 

llebens. Hcbenstreit. 

ihdiv. HtMiwig. 

Ilegelm. Hegel maicr. 

llegetsch. Hegetschweller. 

Heist. Heister. 

lleldr. Heldreich. 

Helic. Helwing. 

llemsl. Hemsley. 

Henck. HenckeL 

Hen/r. Henfrey. 

llemd. Henslow. 

Herb. Herbert. 

Herm. Hermann. 

////(/. Hildebrand. 

Hochst. Hochstetter. 

Ilnffm. G. F. Hoffmann. 

//. Hoffm. Hermann Hoffmann 

Hoffmanns. Hoff mannsegg. 


















Wm. J. Hooker. 
J. D. Hooker, son. 

Humboldt, Bonpland, and 

Jacq. = N. J. Jacquin. 

Jacij.f. J. F. Jacquin, son. 
J. St. Hil. Jaume 8t. Hilaire. 

Jord. Jordan. 

Juntjh. Junghuhn. 

Juss. A. L. Jussieu. 

Adr. Juss. Adrien Jussieu, son. 

Kcemp. Ksmpfer. 

Karst. Karsten. 

Kaulf. Kaulfuss. 

Kindb. Kindberg. 

Kirschl. Kirschleger. 

A7^. Kitaibel. 

Kcdr. Koelreuter. 

Korth. Korthals. 

Kostel. Kosteletzky. 

Kremp. Krempelhuber. 

Kromb. Krombholz. 

Kuetz, Kuctzing. 

L. Linnaeus. 

LabiU. La BUlardi^re 

Last. Lsestadius. 

Lag. Lagasca. 

Lall. Lallement [Marck). 

Lam. Lamarck (Monnet de La 

Lamb. Lambert. 

Lamowr. Lamourouz. 

Langsd. Langsdorf. 

La Peyr. La Peyrouse. 

La Pyl. La Pylaie. 

Ltdeb. Ledebour. 

Lehm. Lehmann. 

Lem. Lemaire. 

Lesq. Lesquereux. 

Less. Lessing. 

Lestib. Lestiboudois. 

L^o. Ldveillc'. 

L'Her. L'Heritier. 

Ulltrm. L'Herminier. 

Liebm. Liebmann. 

Ughif. Lightfoot 

Ulij. Lilijeblad. 

Lindb. Lindberg. 

LindU. Lindblom. 

Lindenb. Lindenberg. 

Lindh. Lindheimer. 

Lindl. Lindley. 

Linn. Linnsus. Also L 

Linn.f. C. Linnaeus, son. 

Lodd. Loddiges. 

Latfl. Loefling. 



Lets. = 


Naud. = 

= Naudin. 







Nees or 

I C. F. Nees von Esenbeck 




Lid to. 


T. Ntes 

T.F.L. Nees ron Esenbeck 




















Magnol. [stein. 



M. Bieb. 

Marschall Ton Biebcr- 




Humphrey Marshall. 






. Nylander. 
















Medikus or Medicus. 




1 Mcisner or Meissner. 

D. Olio, 

D. Oliver. 



A. or C. d*Orbigny. 

















P.deBeauc.V&Msoi de BeauToia. 


[ Andre Michaux. 






Michx. f. 

F. A. Michaux, son. 








Philip Miller. 



Mill. J. 

John S. Mueller or Milk r. 












John Mitchell. 






J. E. Planchon. 



G. Planch. Gustavc Planchon. 






C Montagnc. 


Plumier, Lat. Plumerius. 








Pol ret. 













Muell.Arg.J. Mueller of Argau. 



F. Mm II. 

Ferdinand Mueller. 



0. MiuJl. 

Otto Mueller of Denmark. 










J A. Murray. 



A. Marr. 

Andrt»w Murray. 















Reich. = 

= Reichard. 

Scop. — 



H. G. L Keichenbacli. 




/. li. G. Reichenbach, son. 




















L. C. Richard. 




1 Achillc Richard. 







John Richardson. 




















J. J. Roeroer. 



M. J. /2a;m.M. J. Kociiicr. 



Ram. ^ 

1 Roemcr & Schultes. 














Thomas Thomson. 



























Torr.i- Gr.Torrey & A. Gray. 















St. r/ii. 

A. Sain^Hilaire. 







S<i/m-Dyck.Vnnve Jos. Smlm-Riffer- 




Sau8sure. [schied-Dyck. 
















Veillard or VieUlard 












VUlars, or \lllar. 










) C. H. Schultz, Bipontinus 
J (Zweibrucken). 





T. Vogel. 

























WaUr. = 


WUdb. = 

: Wildbrand. 


















P. W. Watson. 



H.C.Wats.ll.C. Watson. 



S. Wats. 

Sereno Watson. 




























yEst. Estate, in summer. 

^'Est. Estivation. 

Alb. Albumen. 

Anth. Anther. 

Art. Artificial. 

Auct., Auctt. Auctorum, of authors. 

Ant. Autumnal. 

B. or Btat. Beatus, " the late," re- 
cently deceased. 

Br. Bract. 

Cal. Calyx. 

CtL Celeberrimus, or Very cele- 

Cent. Ci'ntimetre. 

CI. Clarissimus. 

Char. Character. 

(V>//. Collection. 

Cor. Corolla. 

Cult. Cultivated. 

Deri in. or Dfc"*. Decimetre. 

I>€$cr. Description. 

Diff. Differentiae, the distinguishing 

Ed, Edition. 

Emlrr. Embrvo. 

Esit. Essential, as Char. Ess. 

E.rcl. 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. ^ 

Fa-m. Female plant, flower, &c. 

Fol. Folium, leaf. 

Fr. Fruit. 

Frnctif. Fructification. 

Gen. Genus or Generic. 

Germ, Germen, Linnaean name for 
ovary ; also Germination. 

//. Herbarium. 

Ilab. Habitat, place of growth ; 
sometimes for Habeo, I have. 

F/trb. Herbarium. 

Ifort. Hortus, garden. 

Hortul. Hortulanorum, of the gar- 

Ic. Icon, a plate or figure. 

///. Ilhistris. illustrious. 

Iiifd, Unpublished. 

/«/! Inferior. 

//'//. Inflorescence. 

Inr. Involucre. 

Lat. Lateral, or relating to width. 

Lin. Linea, a line (the I2th of an 

Lit., Lift, In a letter or letters. 

/. c. Loco citato, in the place cited. 



Mtisc. Male plant, flower, &c 

Mill, or mm. Milltnietre. 

Ms$, Manuscripts. 

J/f<5. Museum. 

N. or No. Number. 

Nnt, Natural. 

Nom. Nomen, name. 

Obs. Observation. 

Ord. Order. 

Oo. Ovary. 

p. Page, or sometimes Part. 

Ped. Peduncle or Pedicel, or for 

Pedalis, a foot long or high. 
Peric. Pericarp. 
Perig. Perigonium. 
Pet, Petal or Petiole. 
Pist. Pistil. 
Plac. Placenta. 
Poll. Pollicaris, an inch long. 
p. p. Pro parte, in part. 
Prodr. or Prex/. Prodromus. 
Rad. Radix, root ; or Radical. 
Ram, Ramus, branch. 
8, Seu, or Sive, Latin for or. 
Sect. Section. 
Segm, Segment. 
Stm. Semen, seed. 

Sep. Sepal. 

6V. 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 7a6. Tabula, plate. 

T. Tomus, volume. 

V. Volume : sometimes for Vel, or; 
sometimes Vide, see. 

Var, Variety. 

Veg, Vegetation, characters of 

Vern, Vernal. 

t;. 8, 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- 

V. 8, c. and v. a. s., indicates that 
the dried specimen was cultivated 
(c) or spontaneous («). 

V. V, c. and r. v, s., that the living 
plant seen was cultivated (c) or 
spontaneous («). 



O An annual plant. 

$ A biennial. 

11 A perennial, 

b A tree or shrub. 

* Affixed to a reference, means that a good description will be found there, 

t Indicates an obscure or doubtful species. 



O A monocarpic plant, i. e. which dies after once flowering and fruiting, 

either annual or biennial, or of longer duration. 
vL) Annual. 
(?) Biennial. 
@ Monocarpic perennial, such as Agave. 

892 siOKa. 

y. Perennial herb. 

^ SuffruifT, an undcrthruK 

*) Frtitex, a shrub. 

5 Arbuicula, a tree4ike shrub of teo to twentj-flw fctt ui htt^ki 

5 Arbor, a tree. 

r^ A climbing plant 

A An evirrgrevn. 

^ Male plant or flower. 

9 Female plant or flower. 

^ Hernia plirodite plant or flower. 

00 Indefinitely numerous, t. g. co-nndra, pol/amlroas. 

? A sign of doubt. " Thalictmm 7 Jtiptmimm'* douUs if tW fiam* m 
really a Tlialictrmu. ** Thatirtrmm Jitftomirwm, Tbuab ' " ^mku 4 -tm 
pUnt in liand is truly tlie sitecK-s of Thunbrrg JlLm^rtrum Jt,mm 
cum, Thunb.. WiUd.? doubu wbetbtr Wilidefiow's T J^fmm.',m « 
really tliat of Tliunberg. 

1 A sign of certainty. As " Tkalirlrmm amfmamtdr; Hirht * Fl frir A« 
p. 822,** as used by lleCandoUt, aflrms that he has srrs an sttiW«t« 
original specimen of tliis author. Afllxed to the naosr ol a fT^;ierv# 
as ** Virginia, Ciai/tom ! " it afllmis that the writer has •lai 
specimen collected by the person to whose name it is apprttiM. 

- Between two flgures, as In " Sumens 6-10.** indicates the •at 

difference, as that the stamens are from flve to ten. 
^ ' " The signs for degrees, minutes, and seconds, as P. 3'. t *. aiv 

Gray's Msnoal of Botany of the Northern United Scales, for tnt '■ 
liuhi'S ('). and lines (") With Kur«i|»esn auiliors. usually tb« • g% 
for minutcn in for tei't ; that <»f seconds fur imhcs thus 1 . s i tA 
hi^rh ; 1", sn inch long ; and 1'", a line long. 
0= (\)tyk'd<)ns Accunil>ent to the radicle. 
Oil Cotyledons incumbent on the radicle. 






This Glossary is intended to contain all the principal technical terms (substan- 
tive as well as adjective) of siructural iCnd systematic Botany, as far at least as 
concerns Phaenogamous plants. Most of the special terms relating to the lower 
Cryptogamia 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; 
ay, ylpetalous, destitute of petals; 
Aphyllous, leafless. In words be- 
ginning with a vowel, this prefix is 
changed to an; as, ylnanthous, flow- 
erless ; ylnantherous, antherless. 

AbbrtvUitiotu^ 38o. 

Aberrant. Wanderinir, applied to spe- 
cies, genera, &c., which differ in some 
respect from the usual or normal char- 
acter of the group they belong to. 

Abnttrmnl {Abnormia). 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 {rivut). Defective or barren. 

Abrupt (Abruptus). Terminating sud- 
denly ; the opposite of tapering. 

Abruptly pinnate. Pinnate without a 
terminal leaflet or appendage ; 101. 

Aeanthdcladout {-ut). Having spiny 

Acanthdpharout (-ui). Spine-bearing. 

Acaulesceni i-tm). Stemless, or appar- 
ently so, with no proper ciu/m ; 45. 

Acaulit. Stemless ; same as Acaulescent. 

Accettory. Something additional, or of 
the nature of appendage. 

Acceuory Budn. 44. 

Accetanry 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 bodv. 

Accumbent Cotyledons. With edges 
againiit the radicle: 313. 

Acephalous {-us). Headless. 




Mtrmt {-6tu$), NeedU-fthapftI, like the 
leave* of Pin«*9. 

Acittihulijorm (-^frmis). In the (urm of 
a hhallow open cup ur »aui-«*r. 

Arh(rniHm or Arktnimm. A Ainall, dry 
aii«l luinl,one-<TU«Ml, uiie-M^irtl, iiul«- 
bi»i'«riit fruit; htru'llv one of a »iii|;le 
and fnt' carfwl; hut vxtirudvil to Mini- 
lar oncn of nmrv than one carprl. and 
aiwuiith adnatt* calyx ; ^A. {Arktg- 
nium i» ctymologii-ally the pro|^rr 
orthof^niphy ; hut achtmium is he- 
mniinfc the commoner form.) 

ArhttntHiirp {tirpium). (ieneral name 
ii( a dry and indehiMrent iruit : il^. 

ArhnnifiiuiH. Such a douhir achenium 
a* that of I'mhellifenr ; a Crrmv- 

AcMLitujftletms {tms). Dctlitute of peri- 
anth; lUl. 

AciruLi. A briktle. 

Arimliir {-arts). Hrtitle-fthapcd, or »leo- 
dcr neetlle-fhaped. 

Acinarijorm {-4trmU). Sc« milar-Miaped ; 
rurvtMl with n>unded |Hiint, thicker on 
the ^traif;hter ed^pe than ou the con- 
vex edge. 

Acimttui. Like p^pe^ or f(ra(ie-»ee<l. 

«irtMM«. ClaMiically a lM>rry, |Mirtiru- 
larly a gra|»e, or iXn ^tony M^*d. or a 
iMinrh (d berries; now Miuieiiuin* ap- 
plicil to the M-|iarate caqirU of an 
ac;;n-^nt«« lia<'rate fruit, or to the ron- 
tainetl M«>n«' or M'e«l : 21>7. 

At>trn. Kruit <»f Ihe (»ak. 

.|(i»/y/<f/<«N, |tl. Afnt^lnittnt, At'"fyl'*l>mrt. 
A plant or plantt dt-Miiute of coty- 
IimIlii. ur 

A*'"tiil"limi u$ (-/iM). Without cotyle- 
«|on«: a* thf fnihrvo of <'uM-uta: *i»», 
^{H Mostly apfilii-d. a* hy Ju*«iru. 
to plants whii h hjv«* lit* pr<i|ifr **r*\ 
ii«ir iiiilir^o, anil thcrffoiv no vij'ylc- 

Ai'i'-niifiKthrtfii. I'laitt* prtnlu* in;: "ide an 

wi!!a«« ttTiiiiiialhuiUor croHth- : •i41 
-d'"'««'" V'- riants ^:rl•win^ from a|n*\ 

oiilv ; :ui. 
,4<-»ii',f n .l''r6t}rntr). Nanif of ria** of 

plants Mlii«h in growth an* «ai<l to In* 
A''' ' rtfUM. (iriiwin^ fr«»ni iIm- a|nx 

or t>v ti nniuiil hu<!« otil\- 
Af i^iri'utn. lK«>uu\'» nanii* for a 

I* rry fn»ni an o\ary with adnale 

.1.- ■■•y.ioi. An olil nam*- «-f thf plu- 

iM'.i!.- (if a crain in ;;«nm nation. 
.!...'# *t, ,.<,itu») I'ri»;WI\ ; l>fM.t with 



ArmlntMti* \^im»y IW««t m^\ 4a« u ^ 

toe prickW*, or 
AfmlnUk. llMuAuln * a4 a> % « 
ArmUm*. A prvkv . a )«•--••: m»a 

exirrttrriKr of lb* t«ark 
.IriiMra. A Uf^ru..: i*^.^? 
.Imwiarirc I -^Ims . I^apiw^ &a a taj** «^ 

poiul ; 1M. 
<iir«f« ( .Im/iuj. Trmiinaiitiig ta m » » ^ 

aui;lr: «7. 
AcntpfUtl. lHt