G S7SZLS10

|

OLNOHOL JO

Pee POTANICAL® SEX EnOOK
Vo ee 1;

SRAUOTHAL ema vy.

Digitized by the Internet Archive
in 2010 with funding from
University of Toronto

http://www.archive.org/details/graysbotanical01 gray

GRAY’S BOTANICAL TEXT-BOOK.

VoLumME I.

STRUCTURAL BOTANY.

GRAY’S BOTANICAL TEXT-BOOK

CONSISTS OF

Vou. I. Srrucrurat Borany. By Asa Gray.
II. PuysioLoeicat Botany. By Grorce L. Goopate.

Ill. Intropuction To Cryprocamic Botany, BOTH
o
StrucTURAL AND Systematic. By Wi iiam G.

Fariow. (Jn preparation.)

IY. Sxetcu or THE NatTuraL OrpDEers 0F PH£NOGAMOUS
Piants; their Special Morphology, Useful Pro-
ducts, &c. (Jn preparation.)

au

GRAY’S BOTANICAL TEXT-BOOK.

(SIXTH EDITION.)

Wor. IE.

SrRUCTURAL . BOTANY

OR

ORGANOGRAPHY ON THE BASIS OF

MOA? TOLOG YY:

TO WHICH IS ADDED THE PRINCIPLES OF

TAXONOMY AND PHYTOGRAPHY,
AND

A Glossary of Botanical Terms.

BY
ASA “GRAY, LL.D., Ere.,

FISHER PROFESSOR OF NATURAL HISTORY (BOTANY) IX
HARVARD UNIVERSITY.

IVISON, BLAKEMAN & COMPANY,

PUBLISHERS,
NEW YORK AND CHICAGO.

Copyright,
By Asa GRAF,
1879.

PREFACE.

THE first edition of this treatise was published in the
year 1842, the fifth in 1857. Each edition has been in
good part rewritten, —the present one entirely so,— and the
compass of the work is now extended. More elementary
works than this, such as the author’s First Lessons in
Botany (which contains all that is necessary to the prac-
tical study of systematic Phznogamous Botany by means
of Manuals and local Floras), are best adapted to the
needs of the young beginner, and of those who do not
intend to study Botany comprehensively and thoroughly.
The present treatise is intended to serve as a text-book for
the higher and completer instruction. To secure the
requisite fulness of treatment of the whole range of sub-
jects, it has been decided to divide the work into distinct
volumes, each a treatise by itself, which may be indepen-
dently used, while the whole will compose a comprehensive
botanical course. This volume, on the Structural and
Morphological Botany of Phznogamous Plants, properly
comes first. It should thoroughly equip a botanist for the
scientific prosecution of Systematic Botany, and furnish
needful preparation to those who proceed to the study of
Vegetable Physiology and Anatomy, and to the wide and
varied department of Cryptogamic Botany.

1v PREFACE.

The preparation of the volume upon Physiological
Botany (Vegetable Histology and Physiology) is assigned
to the author’s colleague, Professor GOODALE.

The Introduction to Cryptogamous Botany, both structu-
ral and systematic, is assigned to his colleague, Professor
FARLOW.

A fourth volume, a sketch of the Natural Orders of
Phenogamous Plants, and of their special Morphology,
Classification, Distribution, Products, &c., will be needed
to complete the series: this the present author may

rather hope than expect himself to draw up.

ASA GRAY.

HERBARIUM OF HARVARD UNIVERSITY,
CamBripGe April 10, 1879.

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

CONTENTS.

; PAGE
Y INTRODUCTION. THe DEPARTMENTS OF THE SCIENCE 1

CHAPTER I. OUTLINES OF THE GENERAL MORPHOLOGY
OSE HE NOGAMOUS ELANTS) = 0s ef 2 21s 6 5

CHAPTER Il. MORPHOLOGY AND DEVELOPMENT OF THE

EAB RVOSAND. SEE DEUNG: 2.0 ee cn) een © 9
The Embryo, its Nature, Structure,and Parts . . ... . 9
Development of the Dicotyledonous Embryo in Maple . . . 10
In Ipomea, or Morning Glory, &c., with Albuminous Seeds . 13
In Embryos with thickened Cotyledons . . . ..... 16
As of Almond, Beech, Bean, &e.. . . . 17

With Hypogeous Germination and no Elongation ‘of Grable 19
In Megarrhiza, &c., with concreted Petioles to the Cotyledons 21
In Ipomea leptophylla with foliaceous and long-petioled

Cotyledons and no elongation of Caulicle. . . . . . . 22

In Pumpkin, &c., with no Primary Root . ....... 2
The Polycotyledonous PMDLY Ones ee 23
The Monocotyledonous Embryo of Iris, Onion; ‘Cereal Gane 24
_ Pseudo-monocotyledonous and Acotyledonous Embryo. . . 26
Dicotyledonous and Monocotyledonous Plants . . . . . - 27

CHAPTER II. MORPHOLOGY AND STRUCTURE OF THE

ORGANS OF THE PLANT IN VEGETATION .. 27
PSECHLON La ORC THE ROOTS ToL: pr lise he spades fuss le Va ea oll PL
Nature, Growth, and Compstition mera tee eh EDS
LSYGTA TERNS Berean Oe ati aes 8 Me ite A ie ale lah ee Mey RI
BITIASFOMMOOLSteeMC nin ons I ncas ee cal ee eee Atos See eee
Dna OncmATTIaIse a cs Eacsa- tu eo eas os to ek ee. 80
ICH DIGae Res ee Subse s Sol APC. FN ts mares OR
IBCKenni ashy wbowe See ty ub) ceca ee oe at eee a Be
Aerial Roots shies Ba ey oie ey et A a eT a es, Ee
Epiphytes or Air- Seilasile aswel. 5 Se Thee ape ks

Parasitic Plants, Green and Colored He Aap ee ee Fe orn Os

CONTENTS.

Section II. Or Buns

Scaly Buds and Bud-seales . :
Naked, Subpetiolar, and Fleshy Buds :
Bud-propagation :
Norinal, Accessory, and desi: Buds :

Section III. Or tHe Stem

§ 1. GeNERAL CHARACTERISTICS AND GROWTH

Development and Structure
Ramification, Branches ‘

Excurrent and Deliquescent Stems :

Definite and Indefinite Annual Growth

§ 2. Forms or STEM AND BRANCHES

Herbs, Shrubs, Trees, Culm, Caudex, Scape
Climbing Stems, Twining or otherwise
Leaf-climbers, Tendril-climbers, and Rept-elarbers
Suckers, Stolons, Offsets, Runners

Tendrils formed of Stems F
Sympodial and Monopodial Stems

Spines or Thorns and Subterranean Stems
Rhizoma or Rootstock

Tuber, Tubercles

Corm or Solid Bulb

Bulb, Bulblets

Condensed Aerial Semen

Stems serving for i Phyllocladia, » Cladophy lla

Frondose Stems .

§ 3. INTERNAL STRUCTURE

Anatomical Elements .

Endogenous Structure

Exogenous Structure ; its Besiaiane

First Year’s Growth

Pith, Layer of Wood, &e.

Bark, its Parts and Structure

Annual Increase in Diameter .

Demarcation of Annual Layers

Sap-wood and Heart-wood .

Growth and Duration of Bark * :
Living Parts of a Tree or Shrub, Longevity .
The Plant composite .

Section IV. Or LEAVES

§ 1. Tuerr NaTuRE AND OFFICE

Parts of a Ileaf .
Duration, Defoliation, Normal Beacon

CONTENTS.
§ 2. TuHerr SrrucTuRE AND Forms As FOLIAGE .

Internal Structure or Anatomy

Parenchyma-cells

Epidermis, Stomata or roving: pores

Framework, Venation . . Sa ees

Parallel-veined or Nerved ee CS er ween Bat So) ne

Reticulated or Netted-veined Leaves

Pinnately or Feather-veined and Palerdtely or adiately
Veined aN ED

Honma as tonOutdiney oye vo es ke (oss es ee Ee

Forms as to Extremity ;

Forms as to Margin or Special G@eadine and Dentekon -

ihobationvor/Seomentation «2 4 2s. | 2) 20's. ae

Number and Arrangement of Parts .

Compound Leaves, Pinnate and Palmate or Dipitate &e:

Petiole or Leafstalk

Stipules, Ligule, Stipels .

Leaves in unusual Modifications .

Such as Inequilateral, Connate, Periolate

Vertical and Equitant

Without distinction of Parts

Stipules serving for Blade

Phyllodia, or Petioles serving for Blade

§ 3. LEAVES SERVING SPECIAL OFFICES

Utilizing Animal Matter
Ascidia or Pitchers
Sensitive Fly-traps

Leaves for Storage :
Bulb-scales and Bud-scales .

CHAPTER IV. PHYLLOTAXY, OR LEAF-ARRANGEMENT

Section I. DistrisuTion or LEAVES ON THE STEM

Phyllotaxy cither Verticillate or Alternate, Cyclical or Spiral
Verticillate or Cyclical Arrangement

Alternate or Spiral Arrangement

Its Modes and Laws :

Relation of Whorls to Spirals .

Hypothesis of the Origin of Both

Fascicled Leaves

Section IJ. Disposition or LEAVES IN THE Bup .
Vernation and Mstivation; the Modes .......4.
Direction. Dextrorse and Sinistrorse .......

Vili CONTENTS.
CHAPTER V. ANTHOTAXY OR INFLORESCENCE... . 141

Bracts and Bractlets and their Modifications . . .. . . I4l

Peduncles, Pedicels, Rhachis, Receptacle ..... . . 143
Position of Flower-buds, Kinds of Inflorescence ... . . 144
Indeterminate, Indefinite, or Botryose. . . ... . . . 146
Raceme, Corymb; Umbel) 2 5) 3) ae) Sec a
Meador Capitultnige-aysee nae Reet Go 1Gt/
Syconium or Hypanthodium ..... . . 9. «. » «= «= 148
Spike, Spadix, Ament or Catkm .©. .-. «= % 205 =e
Panicle and other Compound Forms ....... . . 180
Determinate or Cymose.= -.- @ a) =a
Cyme, Glomerule, &. . . Mera an sc LE
Botryoidal Forms of Cymose Ty oe | sts sta: Sap shea ee ee
Sympodial Forms. . . 154
Scorpioid and Helicoid, the Pinatas oe “4
Monochasium . . PR ESS
Bostryx, Cincinnus, Epadine i omen ‘&e Me re EY
Mixed Inflorescence =<. «i+ 2: By = & 4. tes 6) eens
Thyrsus, Verticillaster,&c. . . a: rb nales) PET
Relations of Bract, Bractlet, and lower “oe eu pt ee
Anterior and Posterior, or Inferior and Superior . . . . . 160
Mediamand: ‘Transverse! = <=) q: «) o<l 6) oscuro
Position:of Bractlets: sn. Gy a a) a te oe ee
Tabular View of Inflorescence .. - . . 2 « % » « =» aioe
CHAPTER VL THE FLOWER . =. ..>< =. =. ee
Section I. Irs Nature, Parts, anpD MeTaAMoRPHY .. . . 163
Floral Envelopes, Perianth, or Perigone . . ..... . 164
‘The Parts, Calyxsandi@orolla, 7 <2) sew 2) eee
Androecium, Stamensy.° . <=. = = -gen. (. 1) aeons
Gynecium; Pistils.-y 3. oe iy a ie Le ee fe eee
Torus or Receptacle of the Flgsres oy! eo joc ta ee
Metamorphosis . . . oe) 3o- OI
Unity of Type cient i Posae oad Fcsuasititners «ee ED
Teratological Transitions and Changes . ...... . 170
Srorion, Dk, Bio ran, SyMweRRY 2. % 2.0 2) <5 2 ele ee ee
Symmetrical, Regular, and Complete Flower . . .. . . 1785
Numerical Ground-plan-.) < : = < =o. ssi, 2) sures
Pattern. Flowers’02 1. 8 0s gen so sae) ee
Diplostemonous Type... <= =... sy ea
Section III]. Various MopiricaTions OF THE FLOWER. . . 179

§1. "ENUMERATION Or THE KINDS -. 2% =<) < =i. seseeeeeee

§ 2. Recurar Union or Srmiuar. Parts ..-. ... . 180

Coalescence or'Cohesionicus 60 )sn0) ame 2. eee) eee eee

CONTENTS. jie.

§ 3. Union oF DisstMILar OR SuccESSIVE Parts . . . . 181
Adnation or Connation . . Cer Re ren oan vated oh LOD
Hypogynous, Perigynous, papers NG Te RCE aah lee iceaar ia tS;

§ 4 I®REGULARITY OF StmmtaR Parts ....... . 184

§ 5. DrisAPPEARANCE OR OBLITERATION OF Parts . . . . 187
Abortion or Suppression of Parts of a Circle . . . . . . 187
Abortion or Suppression of whole Circles. . . . . . . . 190
‘Lemus therewith:conneeted) | Eos .0 so ee eee «(LOL
Suppressed Perianth . .. . Te yc oe eee op BLO
Suppressed Andreecium or Coane Sate: (Akiw cs. gene er OS
Alone with suppressed Pertanthy <5 1. «os ste « = 194
INetira eNO WeCsiu. saath ee commerce cM sls) oso as, ae ante a OD

§ 6. INTERRUPTION OF NORMAL ALTERNATION . ... . . 195
Anteposition or Superposition. . . .... ome Ca mee a eie
eAippaarance only” 285 an Pea ie a ot Be PT 196
Superposition by Spirals ... . Eanes ap LOG
Anteposition with Isostemony and Ds eee aelomrepe Ws LOG
MiEh@bdiplostemony ys). 2. sh Sacete 6. fe oe) OR eae tee 98

Si) INCREASED NUMBER OF PARTS’. «29. 5 2%. » « « 200
Remar vidlimplicaione 6 «she Ge) 4 ep ee ae on 200
iParapetalous: Multiplication <9. oi ¢9 st aes ts ee 2) OL
@horisissorMeduplication.. 2s) - = wares ot a 6202

SSamOULGROWDHS 0 ses ele eso eee, St (eS RM SIE A. ‘DOO
Their relation to Chorisis: -Trichomes. . . .... . . 209
ComonmiomCrowir i203 ces, Bikes soon es aoe ae. . 210
ipl ewe eaes (sec! coe Wea oe ee re ee eles ZIT

§ 9. Forms or THE Torus OR RECEPTACLE. . «. ... . 2i1
Stipe, ea Cae SEER ORES: CHER re One Pe are 1
DISkar rere! : Se Fe 5;
yp am COMOI Nes sateiy a po. Boa. ee ee or ictikrs eon ot LA:

Section IV. ApapraTIoNs OF THE FLOWER TO THE ACT OF
HGRIEGI TA MMTOND casein) ig ttl ft oem ae 2) Lees pete is 2A
Reem eLNN GrENTURVAE 0 nctiecy ius! Bg") s<s--y iota” Rise at uk Waceeieele: .< 2L5

Close and Cross Fertilization, or Autogamy and Allogamy . 216

§ 2. ADAPTATIONS FOR ALLOGAMY OR INTERCROSSING . . . 216
Wind-fertilizable or Anemophilous Flowers. . . . . . . 217
Insect-fertilizable or Entomophilous Flowers .... . . 218

Irregularitv as related to Allogamy ......... 219

CONTENTS.

Dichogamy, either Proterandrous or Proterogynous .
Proterogyny .

Proterandry

Particular Adaptanent in Pea pionaeeoe Flowe ers

In Kalmia-blossoms, Iris, &c.

Transportation of Pollinia .

In Orchidacew and Asclepiadacee

Heterogonous Dieter and menses

§ 38. ADAPTATIONS FOR CLOSE FERTILIZATION .

Cleistogamy .

Section V. THE PERIANTH,. OR THE CALYX AND COROLLA IN

PARTICULAR

Perianth as to Duration, Numerical Terms, Union, &c.
Parts of Petals and of Gamophyllous Perianth .
Forms of Corolla and Calyx ‘

Section VI. Tue ANDR@CIUM, OR STAMENS IN PARTICULAR .

The Stamen as a whole; Numerical Terms . ;
The Filament and the Anther; their Modifications
Pollen

Pollen-tubes .

Section VII. Tue Pistits, or GyN@&CIUM
-§1. IN ANGIOSPERMS .

Carpel or Carpophyll . oe At

Ventral and Dorsal Sutures; Placenta

Simple or Apocarpous Pistils .

Compound or Synearpous Pistil .

With two or more Cells and Axile Piacantee Partitions
With one Cell and Parietal Placente

With one Cell and Free Central Placenta

Anomalous Elacentatiom << "= Welerss Were! bee: ae

§ 2. In GYMNOSPERMS .

Structure in‘Gnetacee . . . = «. =» « «
Simin Commies es 5 9 6 6 o o - o 9 O-¢
In the Yew Family

In the Pine Tribe, &e.

In the Cypress Tribe .

Structure in Cycadacee .

Section VIII. Tue OvuLe

Its Structure and Position .

Its Forms, Orthotropous, Campylotropous, ee

Anatropous
Origin and Morgholamenl aan ae pas Omles
Origination of the Embryo .

CONTENTS.

CHAPTER VII. THE FRUIT

Secrion I. Irs Srructure, TRANSFORMATIONS, AND DEHIs-
CENCE.

Pericarp, its Alterations, Accessions, and Transformations
Dehiscence

Section II. Tue Kinps or Fruit

Simple Fruits : jth

Dehiscent Fruits, Follicle, Tenn. eaeaule iy xis Signe

Indehiscent Dry Fruits, Samara, Akene, oe Caryopsis,
Nut, &e. . .

Fleshy Fruits, Drupe, pea pene ery. &e.

Aggregate Fruits Se heh

Accessory or Anthocarpous ineiies

Multiple or Collective Fruits, Syconium, Strobile. &e.

Table of Simple Fruits

‘CHAPTER VIII. THE SERED

Its Stalk, Coats, and ae
Aril or Arillus :

Nucleus or Kernel, Albumen

The Embryo, its Parts and Positions

The Cotyledons as to Adjustment and Sen:

CHAPTER IX. TAXONOMY .

Section I. THe PRINCIPLES OF CLASSIFICATION IN NATURAL
History

Individuals

Species .

Varieties, iaaece: digs

Cross-breeds and Hybrids

Genera . ; :

Orders, Classes, Tribes, Ber

Sequence of the Grades . one

Nature and Meaning of Affinity . i
Theory of Descent and Natural Selection

Section II. Boranicat CLassiFICATION

Ante-Linnzan Classifications .
Linnean Classification

Sexual Artificial bee
Natural System

As presented by esi ;
Some of its Modifications

xi CONTENTS.

CHAPTER: X, PHYTOGRAPEY 5 cic. eee cyt eee
Section 1. NOMENCLARURE” .4.)5 <5 *. 12) «0» 5h see eS
Names of Plants, Binomial Nomenclature ..... . . 3846
Rules fornaming:; Plants “is... = i cy cyte eee
Names of Genera . . . PO ce ane errs
Names of Species, Neeretien! on hotene ee en 50)
The Fixation, Precision, and Citation of Gavies aE
Subgeneric Names 2 5.9". 7 4 2) 58
Tribal'and Ordinal’Names'. ~ 2 2% 4° .° 20a) 2 cee
Names of Cohorts@lasses. Ge 35:5). ee EES
Section II. GLossoLoGY OR TERMINOLOGY .-..... «. ooo
Secrion JIL. DrscRIPTION.© : . 2956 6 6) 2 6 eon
@haracters- 4.03. Ss oe ee ee, ne Ca
Punctuation . "4 6 ee ee eer Se
Synonomy «2 o eee oe ee
Iconography. . . . © Be ee, G os 2a noe
Habitat and Station, ee alee Dat PRL al er, ec
Rtiymology of Names: =. 297 2) =) cue che cup Cc eerememmneLEee
Accentuation, Abbreviations 4 .: . « =. . «| =» 0 thenneOOM
Signs .. =. ee wr) we S| chat Sites aon um
Floras, Marcie aoe ee res St
Section IV. Specimens, D1irRECTIONS FOR THEIR EXAMINA-
TION, PRESERVATION, GG) 9: « ‘:\ 2 | & [a = eeeomemCMD
Implements of Investigation . . . . =. 5 = = |) sesneeene
Diagrams...) 32+ -<p .b eof os ws ce gs
Wer bOrizing) ss capers ss i 0 ee ge yeel eee et eee
Drying Specimens.) 9. «cK ee
Poisoning Specimens... 2. so. i ok se oe
The Herbarium... ofS 2%. 266 oe 2S <b eee
ABBREVIATIONS “he 6 st cs eae cee oe oy Ge San SCD
SIGNS. 2.20 )..2 2 4.2.3 904-5 © 204 7 See

GLOSSARY OF BOTANICAL TERMS, WITH INDEX . . 393

STRUCTURAL BOTANY

THE BASIS OF MORPHOLOGY.

INTRODUCTION.

1. Tue two Biological Sciences,’ considered as parts of Natural
History, are Zoology and Botany. The latter is the natural
history of the Vegetable Kingdom. It embraces every scientific
inquiry that can be made respecting plants, their nature, their
kinds, the laws which govern them, and the part they play in
the general economy of the world.

2. We cannot distinguish the vegetable from the animal king-
dom by any complete and precise definition. Although ordinary
observation of their usual representatives may discern little that
is common to the two, yet there are many simple forms of life
which hardly rise high enough in the scale of being to rank dis-
tinctively either as plant or animal; there are undoubted plants
possessing faculties which are generally deemed characteristic of
animals ; and some plants of the highest grade share in these
endowments. But in general there is a marked contrast between
animal ‘and vegetable life, and in the part which animals and
plants respectively play in nature.

3. Plants only are nourished upon mineral matter, upon earth
and air. It is their peculiar office to appropriate mineral mate-
rials and to organize them into a structure in which life is mani-
fested, — into a structure which is therefore called organic. So
the material fitted for such structure, and of which the bodies

1 Biology, the science of life, or rather of living things, in its earlier use
was equivalent to physiology: recently, it has come to denote the natural
history of plants and animals, 7. e. of the two organic kingdoms, including
both their physiology and descriptive natural history.

1

2 INTRODUCTION.

of plants and animais are composed, is called organic matter.
Animals appropriate and live upon this, but have not the power
of producing it. ‘So the vegetable kingdom stands between the
mineral and the animal; and its function is to convert materials
of the one into food for the other. Although plants alone are
capable of building up living structure out of mineral mate-
rials, and are the sole producers of the organic matter which
is essential to animal life, and although animals consume that
which plants produce, yet plants also consume organic matter,
more or less, acting in this respect like animals in all their opera-
tions, except in the grand and peculiar one by which they
assimilate mineral matter. Most plants of the higher grades
assimilate largely and consume little, except in special opera-
tions. Some, on the contrary, are mainly consumers, and feed
upon formed organic matter, living in this respect after the
manner of animals. The living substance of plants and animals
is essentially the same.

4. Botany deals with plants: 1. As individuals, and in respect
to their structure and functions. 2. In their kinds, and as
respects their classification, nomenclature, &c. Accordingly,
the most comprehensive division of the science is into Puysio-
LOGICAL or BroLocicaL Borany (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 Physiology.

5. SrructuraAL Botany comprehends all inquiries into the
structure, the parts, and the organic composition of vegetables.
This is termed OrGANOGRAPHY, when it considers the organs or
obvious parts of which plants are made up, and MorpHo.oey,
when the study proceeds on the idea of type. The term
OrGANOGENY has been applied to the study of the nascent
organs and their development; Puyroromy, or VEGETABLE
Anatomy, to that of the minute structure of vegetables as re-
vealed by the microscope, 7. €. to the composition of the organs
themselves. But, since anatomy in the animal kingdom includes
the consideration of general as well as of minute structure, and
indeed answers to organography, the minute anatomy of both
kingdoms takes the special name of Hisrotocy. The study of
functions, or of the living being (animal or plant) in action,
is the province of PuysroLocy.

6. Systematic Borany, or the study of plants in their kinds
and in regard to their relationships, comprises Taxonomy, or the
principles of classification, as derived from the facts and ideas

INTRODUCTION. 3

upon which species, genera, &¢c., rest; CLASSIFICATION or the
SysTEM oF Piants, the actual arrangement of known plants in
systematic order according to their relationships ; PHyroGrapny,
the rules and methods of describing plants ; and NoMENCLATURE,
the methods and rules adopted for the formation of botanical
names. GuLossoLocy or TrerMrINoLocy’ is a necessary part of
Phytography or Descriptive Botany, and hardly less so of
Structural Botany: it relates to the application of distinctive
terms or names to the several organs or parts of plants, and to
their numberless modifications of form, &c. This requires a
copious vocabulary of well-defined technical terms, by the use of
which the botanist is able to describe the objects of his study
with a precision and brevity not otherwise attainable. It will
be convenient to exemplify the principal terms along with the
modifications of conformation which they designate; and also,
for greater fulness and facility of reference, to append to this
volume an alphabetical summary of them, or Vocabulary of
Botanical Terms.’

7. The present volume is mainly devoted to Morphological
Botany; that is, to Structural Botany on the basis of mor-
phology. This department cannot be properly dealt with apart
from considerable reference to intimate structure, development,
and function, the subject-matter of vegetable histology and
physiology. But these will here be treated only in the most
general or incidental and elementary way, and only so far as
is necessary to the understanding of the morphology of the
stem, leaves, &c. The whole discussion of the histology and
physiology of plants is relegated to a following volume and to
another hand.

8. The most comprehensive and important division of the
vegetable kingdom is into plants of the higher and of the lower
series or grade, 7.e. into PH2xNoGaMmous (or PHANEROGAMOUS) or
FLOWERING, and CryproGamous or FLoweriess Piants. The
first are all manifestly of one type. and therefore have a consist-
ent and simple morphology. The second differ among them-
selves almost as widely as they do from the higher series; and

1 Grossotocy is the better word, but Terminoxoey, although a hybrid
of Latin and Greek, is in common use.

2 What is called GeocrapuicaL Botany is the study of plants in respect
to their natural distribution at the present time over the earth’s surface, and
the causes of it. Fossrr Borany (Vegetable Paleontology) relates to the
plants of former ages, as more or less made known in their fossil remains.
Mepicat Borany, AcricuLtuRAL Borany, and the like, are applications
of Botany to medicine, agriculture, &c.

4 INTRODUCTION.

their morphology is more special and difficult. Wherefore it is
better to treat them separately and subsequently. This will be
done in a third part, by an associate devoted to Cryptogamic
Botany.

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

CHAPTER, E

OUTLINES OF THE GENERAL MORPHOLOGY OF
PHA NOGAMOUS PLANTS.

10. Morrpnoroey, the doctrine of forms, as the name denotes,
is used in natural history in nearly the same sense as the older
term Comparative Anatomy. If it were concerned merely with
the description and classification of shapes and modifications,
it would amount to little more than elossology 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 constructed upon one type
(or ground plan), which culminates or has its archetype in man,
so all plants of the higher grade (8) are strictly of one type;
the different kinds being patterns or repetitions of it, with varia-
tions. The vegetable kingdom, however, does not culminate in
an archetype or highest representative. As respects the organs
of vegetation, the higher classes of cryptogamous plants exhibit
this same type; but it is only in the most general or in a
recondite sense that this can be said of their organs of repro-
duction, and of the less differentiated structure of the lowest
classes. Wherefore cryptogamous plants are left out of the
present view, to be treated apart.

12. Viewed morphologically and as to its component organs,
a plant is seen to consist of an axis or stem, which sends off
roots into the soil, and bears lateral appendages, commonly as
leaves, but which may be very unlike leaves in whole appearance

1 The term Morphology was introduced into science by Geethe, at least as
early as the year 1817 (Zur Naturwissenschaft iiberhaupt, besonders zur
Morphologie, Stuttgart und Tiibingen, 1817-24). On page 9 of the first
volume, he is understood to have suggested this word for the purpose and in
the sense now adopted in botany and zoology. It essentially replaces an
earlier and somewhat misleading word, A/etamorphosis. (304.)

Apparently the first botanist to adopt the term was Auguste de St.
Hilaire, in his “Lecons de Botanique, comprenant principalement la Mor-
phologie Végétale, etc., Paris, 1841. The term seems not to have been taken
up, in zoology, by Etienne Geoffroy Saint-Hilaire, the antagonist of Cuvier
(who was of a wholly different family from that of the botanist), although
the same idea was denoted by his phrase “ unity of organic composition ”

6 GENERAL MORPHOLOGY

and function. ‘These appendages, whatever their form or use,
accord with leaves in mode of origin, position, and arrangement
on the axis or stem. Their most general and ordinary form is
the familiar one of foliage ; hence the name of leaves has been
by botanists extended in a generic way from the green expan-
sions which constitute foliage to other forms under which such
appendages occur. The proper morphological expression is,
that the latter are homologous with leaves, or are the homologues
of leaves.

13. Leaves are borne upon the stem at definite places, which
are termed Nopes. A node may bear a single leaf or a greater
number. When it bears two, they occupy opposite sides of
the stem. When three, four, or more, they divide the cireum-
ference of the stem equally, forming a circle, technically a
Wouort, or in Latin form a Verticit. When only two, the pair
evidently answers to the simplest kind of whorl. So that leaves
are either single on the nodes, in which case they are alter-
nate, that is, come one after another on the stem; or in whorls
(whorled, verticillate), in the commoner case of a single pair
being called opposite. The bare space between two successive
nodes is an IntERNOpDE. This is longer or shorter, according to
the amount of longitudinal growth, which thus spaces the leaves,
or, whorls of leaves, in most various degrees, either widely when
the internodes are elongated, or slightly when they remain very
short. The plant, therefore (roots excepted), is made up of a
series of similar parts, 7. e. of portions of stem, definitely bearing
leaves, each portion developed from the apex of the preceding
one. This constitutes a simple-stemmed plant.

14. Branching is the production of new stems from the older
or parent stem. These normally appear in the Axms of leaves,
that is, in the upper angle which the leaf forms with the stem, —
from which they grow much as the primary stem grew from the
seed. The primary stem, connected with the ground, produces
roots which develop downwardly into the soil, from which they
draw sustenance. Branches, when developed above ground,

1 A common designation for all these appendages being desirable, a good
one is furnished by the Greek name for leaf, @vAAov, PHyLiuM, plural
Pura. This, used with prefixes, may be made to designate the kind of
leaves in many cases, —as, prophylla, cataphylla, hypsophylla.

Recent German botanists use the word Phyl/ome in this sense. It is a
rather convenient and well-sounding word; but phyl/oma 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 phy//um, readily take prefixes, as above, or the adjec-
tive form, as it readily does in prophyllous, hypsophyllous, gamophyllous, &e.

OF PHASNOGAMOUS PLANTS. T
being in organic connection with their parent stem, do not
usually produce roots; but when placed in equally favorable
conditions for it, 7. e. on or in the soil, they may strike root as
freely as does the original stem.

15. An incipient stem or branch, with its rudimentary leaves,
isa Bup. 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- Ne 2e yes
tions, or in a like sense progeny, each of the VY By
preceding one from which it grew. The g YW
simple-stemmed plant is made up of a series

of such growths, each from the summit of r
its predecessor; the branched plant, of ad-
ditional series, laterally developed, from ax-
illary buds. These ultimate similar parts into
which a plant may thus be analyzed, and
which are endowed with or may produce all
the fundamental organs of vegetation, were by
Gaudichaud called Puytoxs. 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 PuyrOmera (qgetor, plant, |
usgoos, part), equivalent to plant-parts, — the
structures which, produced in a series, make
up a plant of the higher grade. In English,
the singular may be shortened to PH¥romEr.
17. 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, as in Acrogenous
Cryptogams, is supposed by some to occur, even normally, in some Pheno-
gams, especially in certain forms of inflorescence; but this has never been
convincingly made out.

FIG. 1. Diagram of a simple-stemmed plant, exhibiting the similar parts, or
phytomera, a to h, of which it is composed.

8 GENERAL MORPHOLOGY

the leaves are alternate; in other words, each phytomer is
single-leaved; while in the subsequent illustrations of plants
developed from the seed, at least the earliest phytomera are
two-leaved.

18. The plan thus exhibited in the leafy stem begins in the
embryo, or initial plant in the seed, and is carried on into the
flower, in which the normal development of the axis finally ends.
One plan prevails throughout. To illustrate it, the morphology
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.

OF THE EMBRYO AND SEEDLING. i)

CHAPTER II.

MORPHOLOGY AND DEVELOPMENT OF THE EMBRYO AND
SEEDLING.

19. The Embryo is the initial plant, originated in the seed.?
In some seeds it is so simple and rudimentary as to have no
visible distinction of parts: in others, these parts may have
assumed forms which disguise their proper character. But every
well-developed embryo essentially 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
normally to be produced. ‘This stem is the primitive internode
of the plant: its leaf or pair of leaves is that of the first node.
The plant therefore begins as a single phytomer. Some embryos
are no more than this, even when they have completed their
proper germination: others have taken a further development
in the seed itself, and exhibit the rudiments of one or more fol-
lowing phytomera. The embryo of the Maple is an example of
the first kind; and, being large 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 embryo
(seen in Fig. 2 in the coiled |
condition which it occupies ©
in the seed, and in Fig. 5
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 seed contains a single embryo. Polyembry, the formation of
two or more embryos, occurs occasionally as a kind of superfcetation in
some seeds. In those of the cultivated Orange it is most common, and an
evident monstrosity. -In Conifer and Loranthaceex, two or three embryos,
of equal size and perfection, are not rarely produced.

FIG. 2. Embryo of Sugar Maple, in vertical section, as coiled in the seed, merely

somewhat loosened. 3. Embryo of same, just beginning to unfold in germination.
4. Same more advanced: a. its stem or caulicle; 0d. its two leaves o» cotyledons.

10 MORPHOLOGY

20. Caulicle or Radicle, and Cotyledons. The name of radicle
was early applied to the axis of the embryo 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 root (24,
44, 78), but a body of the exact nature of stem, from the
naked end of which the root is developed. Wherefore Caulicle
(Lat. cauliculus, diminutive of caulis, stem) is
the appropriate name; and it would be gen-
erally adopted, were it not that the older term
is so incorporated into the language of sys-
tematic botany (in which fixity and uniformity
are of the utmost importance) that it is not
easily displaced. It may be continued in
descriptive botany on this account, but in
morphology it is apt to mislead ; and the name
of caulicle, suggestive of the true nature of
the or%an, is preferable.t The more fanciful
name of Cotylédons was very early applied to
what are now recognized as answering to the
leaves of the embryo: it has the negative merit
of suggesting no misleading analogy.’

21. Development of the Dicotyledonous Em-
bryo, 7. e. the two-leaved embryo. This, in
the Red Maple (Figs. 5-8), usually germinates
in summer, shortly after the fruits of the season
have matured and fallen to the ground. It
differs from that of Sugar Maple in the crump-
i H ling instead of coiling of the cotyledons in the
' seed. Referring the whole physiology of ger-
mination to that part of the work which treats
of Vegetable Physiology, the development of
the embryo into the seedling may here be described, taking that of
a Maple for a convenient type or pattern, with which other forms

1 Linneus called it Rostellum, a name which, being etymologically mean-
ingless in this connection, is not misleading. The French botanists named it
Tigelle, 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 Radicula originated with Gaertner.

2 The name Cotyledon, which was adopted by Linneus, 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 winged fruits of Red Maple (Acer rubrum), with body
divided, to show the seed. 6. Seed extracted and divided, to show the embryo within.
vt. Embryo partly unfolded. 8. Embryo in early stage of germination.

OF THE EMBRYO AND SEEDLING. Fk

may 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 carry the budding end out of the soil and
into the air, where the growing cotyledons 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 lie pointing downward and the root-end
upward in the ground when germination begins, both will curve
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 tendency 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 they are called,
7.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 embryo in
the seed, and until after the full develop-
ment of its cotyledons or seed-leaves,
shows no rudiments of the subsequent
growth. The embryo grows into the plant-
let wholly by the appropriation of prepared
nourishing matter which was provided by
the mother-plant and stored in the seed, —in the case of the
Maple, wholly in the embryo itself, mainly in its cotyledons.

FIG. 9. Maple embryo developed into plantlet of one phytomer, and producing
Tudiments of the second: the lower portion covered with root-airs is the root; the
naked portion above is the canlicle.

12 MORPHOLOGY

After this is consumed and in good part converted into struc-
ture, the plantlet must by the action of its root and leaves imbibe
from the soil and air appropriate materials, and
assimilate them into nourishing matter needful
for further growth. Only then does the rudi-
ment of new structure appear, in the form of a
growing point, or bud, at the node or apex of
the primitive stemlet, between the two seed-
leaves. In this case it soon shows itself as
a second pair of leaves, at first resting on
the node (Fig. 9), next as somewhat upraised
by the development of the second internode
(Fig. 10, summit), and finally both this inter-
node and the pair of leaves complete their
growth (Fig. 11). Then the terminal bud
which crowns the second node develops in
the same way the third pair of leaves and
their supporting internode or joint of stem
(Fig. 12) ; and so on. :

24. The root and the stem grow not only
in opposite directions, but in a different mode.
The primordial stem, pre-existing in the seed
(though at first it may be extremely short)
grows throughout its whole length, but most
in its upper part, so that it may become a
stemlet two or three inches long. But, soon
attaining its full growth as to length, the
stem is carried upwards by the subsequent joints or portions,
similarly developed and elongated, one after the other. Not
that each portion necessarily waits until the growth of its prede-
cessor is complete, — though this occurs at first in seedling Maples
and other embryos unprovided with much store of food, —yet the
development follows this course and order of succession. The
root, on the contrary, cannot be said to pre-exist in the seed, or
at most it may be said to exist potentially in tissue of the caulicle
from which a root or roots normally originate.’ It is formed

ul

1 Yet from nothing which is special to this part of the embryo, nor to the
embryo at all. The primary root is developed from subjacent tissue of the
tip of the caulicle, just as it is sometimes developed from along the sides,
and as secondary roots are from all or most stems under favoring conditions.
This complete similarity, and the fact of what is called the “endogenous ”
origin of roots (i.e. their springing from subjacent rather than superficial
tissue) appear fully to warrant the statement in the text above.

FIG. 10. Maple plantlet with second internode developing. 11. Same with second
internode and pair of leaves complete, and bud of the third apparent.

OF THE EMBRYO AND SEEDLING. a kes)

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

of pre-existing stem, so it goes on to
grow in length wholly or mainly by a
continuation of this formation, the new
at the end of the old. ‘That is, the
root elongates by continual minute
increment of its apex or near it, the
formed parts very soon ceasing to
lengthen. This is in marked distine-
tion from stem, which grows by suc-
cessive individualized portions; and
these portions (internodes), at first
very short, attain or are capable of
attaining a considerable and sometimes
very great, but definitely terminable
length, by interstitial growth through-
out. Moreover, roots are naked, not
producing as they grow either leaves
or any organs homologous with leaves.
They commonly branch or divide, but
in a vague manner; and their new parts bear what are called
Root-hairs, which greatly increase the absorbing surface ; other-
wise they are destitute of appendages or organs.

25. With the Maple embryo, here taken as a type, that of
Morning Glory, Ipomea purpurea, 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 apex. They lie
in contact in Fig. 14, and are very thin, leaf-like, and green
while contained in the seed. Their thinness is shown in Fig. 13,
where a section of the crumpled and folded”embryo, as it lies in
the seed, exactly divides them (passing through the terminal and
basal notches) and also the caulicle, which here is thicker than
both. The germination is similar to that of the Maple; and like
that (as Fig. 16 shows), and for the same reason, no bud or
rudiment of the further growth pre-exists in the embryo or

12

FIG. 12. Red Maple seedling, with three joints of stem and pairs of leaves developed,
the first being the cotyledons.

’

14 MORPHOLOGY

appears in the young plantlet, until that has established itself
and had time to elaborate proper material therefor. This con-
dition is correlated with thin foliaceous
A BS cotyledons, holding no store of nourish-
*/ ment. Here they do not contain sufficient
material for the development of the initial
stem and root. The maternal provision
for this is here stored up in the seed
around but not within the embryo. This
nourishing deposit, seen in the section
(Fig. 15) filling the whole space between
the seed-coats and the thin embryo, was
named by the early botanists and vege-
table anatomists the ALBUMEN of the seed.*
This substance, softened in..germination
and by chemical changes rendered soluble,
is gradually absorbed by the cotyledons
as material for their growth and that of
the developing primary stem and root.

26. Seeds in this regard are accordingly
distinguished into albuminous and exal-
buminous, those supplied with and those
destitute of albumen. The difference
inheres neither in the character nor in
the amount of the maternal provision for the development of
the embryo-plant, but merely in the storage. In exalbuminous
seeds the nourishment supplied for this purpose is taken into
the embryo itself, mostly into the cotyledons, during the growth
and before the maturity of the seed. In albuminous seeds
this same material is deposited around or at least external to
the embryo.

27. The amount of this deposit is, in the main, inversely pro-

1 Grew appears to have first applied this name, and Gaertner to have
introduced it into systematic botany, where it remains in use, although
Jussieu replaced it by the term Perisperm, and Richard by Endosperm,
neither of them much better etymologically than the old word Albumen.
But it must be kept in mind that it was intended to liken the “albumen” of
the seed with the albumen or white of an egg as a body or mass, and not as
a chemical substance; the embryo being fancifully conceived to be analo-
gous to the yolk of the egg, the surrounding substance of this kind not
unnaturally took the name of the white, viz. albumen.

FIG. 13. Section of seed of common Morning Glory, Ipomcea purpurea, dividing
the contained embryo through the centre. 14. Embryo of same, detached and straight-
ened. 15. Embryo in germination; the cotyledons only partly detached from the coat
of the seed. 16. Same, later and more developed, the cotyledons unfolded and out-
spread as the first pair of leaves.

OF THE EMBRYO AND SEEDLING. 15

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 embryo left in

2 al 19 7

™

white ; the albumen 3

24 22 20
as a dotted surface.

The lower rank shows the einbryos detached. That of Mirabilis
has very 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 cotyledons narrowed down to
semi-cylindrical bodies, not leaf-like in appearance, and the two
together not thicker than the caulicle. In Barberry the embryo
is straight, in the axis of the albumen, which it almost equals in
length ; the cotyledons considerably broader than the caulicle,
but short and thickish. That of the Peony is similar, but very
much smaller, occupying a small space at one end of the albu-
men, and seemingly without distinction of parts, but under the
microscope and with some manipulation the broader end is
found to be divided, that is, to consist of two minute cotyledons.
The 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 they develop
in germination.

28. The study of the formation of the embryo in the seed
teaches that all embryos begin with a still more simple, minute,
and homogeneous structure; and these comparisons suffice to
show that all such differences are referable to different degrees
and somewhat different modes of the development of the embryo
while yet in the seed. It also appears that the size and shape

FIG. 17. Section of seed and contained embryo of Mirabilis (IFour-o-clock),
18. Embryo detached entire.

FIG. 19. Section of a Potato-seed. 20. Embryo detached entire.

FIG. 21. Section of Barberry-seed. 22. Embryo detached entire.

FIG. 23. Section of Peony-seed. 24. Embryo detached entire.

16 MORPHOLOGY

of an organ do not indicate its nature, either in the embryo or.
in subsequent growth. But in all the cases yet mentioned the
cotyledons actually demonstrate their
nature by developing in germination
in a foliaceous manner and becoming
the first leaves of the seedling. Nor
is this nature much disguised by the
fact that they differ greatly in form in
different species, and that the seed-
leaves, or developed cotyledons, differ
‘“. much in shape and often in texture
from the succeeding leaves. (See Fig.
11, 12, 25, &c.)

29. To complete the comparison
between the seedling Morning Glory
and that of the Maple, it is to be
noted that here, while the cotyledons
or seed-leaves are two, the following
internode bears only one leaf (Fig. 25), as also will the just de-
veloping third internode; and this continues throughout up to
the blossom: that is, the leaves subsequent
to the cotyledons are not opposite as in the
Maple, but alternate. (13.)

30. All the preceding illustrations are from
embryos which previous to germination have
developed nothing beyond the cotyledons. In
the following, a rudiment of further growth,

8 cre

or a primary terminal bud, is visible in the seed. It is most
manifest in large and strong embryos with thick or fleshy cotyle-

FIG. 25. Further development of Morning Glory, Fig. 16, the root cut away, the
internode above the cotyledons and its leaf completed, the next internode and its leaf
appearing.

FIG. 26. Embryo (kernel) of the Almond. 27. Same, with one cotyledon removed,
to show the plumule, a.

FIG. 28. Section of an Apple-seed, magnified, cutting through the thickness of the
cotyledons. 29. Embryo of the same, extracted entire, the cotyledons a little separated.

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

OF THE EMBRYO AND SEEDLING. LT

dons, ?. e. cotyledons well charged with nourishing matter. The
early vegetable physiologists gave to it the name of PLUMULE
(Lat. plumula, a little plume). The
name was suggested by its appearance
in such an embryo as that of the bean
(Phaseolus), in which it evidently con-
sists of a rudimentary pair of leaves,
while in the pea and the acorn itis a
rudimentary stem, the leaves of which
appear only later, when germination
has considerably advanced. In any
case, the plumule is the bud of the
ascending axis already discernible in
the seed. Fig. 27, a, shows it in the
almond, one cotyledon being removed.
Fig. 28 shows it in the section of a
similar although much smaller embryo,
that of an apple-seed, enlarged to
nearly the size of the other. It is
equally visible in the cherry, the bean,
and the beechnut. The embryo in all
these cases constitutes the whole kernel
of the seed. For the nourishment,
which in all the foregoing illustrations
except the first (2.e. in Fig. 13, 17-23),
is deposited around or exterior to the
embryo, is in these stored within it.
31. The development of these em-
bryos in germination proceeds in the
normal manner, but with two cor-
related peculiarities. First, by the
lengthening of the radicle more or less, their thick cotyledons
are usually raised to or above the surface of the soil; they
expand, assume the green color needful to foliage; but they
imperfectly or in a small degree perform the function of
green leaves. Their main office is to supply the other growing
parts with the prepared nourishment which they abundantly
contain. Then, being thus copiously nourished, the root below
and the ready-formed plumule above grow rapidly and strongly,
haying accumulated capital to draw upon; and the leaves of the

FIG. 31. Beechnut cut across, filled by the fleshy embryo; the thick cotyledons
partly enfolding each other. 32. Embryo of the same in early germination 33. Same
more advanced; the plumule, which is just emerging in the preceding, here developed
into a long internode and a pair of leaves.

2

18 MORPHOLOGY

latter are practically the earliest efficient foliage of the plantlet.
Thus, as in the germinating Cherry-seed (Fig. 30), three or four
internodes of stem, with their leaves, may be produced before
these leaves themselves are sufficiently developed to make any
sensible contribution to this growth. And in the Beech and Bean,

the leaves of the plumule come forward almost before the root
has attached the plantlet to the soil. (Fig. 32, 35.) Between
such cases and that of Maple and the like there are all degrees.
There are also familiar cases in which the storage of nourishment
in the cotyledons is carried to a maximum, with results which
gravely affect the development.

FIG. 34. The embryo (the whole kernel) of the Bean. 35. Same early in germi-
nation; the thick cotyledons expanding and showing the plumule. 36. Same, more
advanced in germination; the plumule developed into an internode of stem bearing a
pair of leaves.

FIG. 37. Embryo of Pea, i. e. a pea minus the seed-coat. 38. Advanced germi-
natior of the same.

OF THE EMBRYO AND SEEDLING.

19

32. Thus, in the Pea, near relative of the Bean, the embryo
(Fig. 37), which is the whole kernel of the seed, has the

cotyledons so gorged with this nutritive
store that they are hemispherical; and
the acorn of the Oak (Fig. 39), near
relative of the’ Beech, is in similar case.
These extremely obese cotyledons have
not only lost all likeness to leaves, but all
power of fulfilling the office of foliage,
which is apparently no disadvantage ; for
when two different duties are performed
by the sime organ, it rarely performs both
equally well. Here they become mere
receptacles of prepared food, the nature
and office of which is the same as of the
albumen, or nutritive deposit exterior to
the embryo in what are called albuminous
seeds. (25-27.) The difference is in the
place rather than in the character of the
deposit. The plumule in such cases is
always apparent before germination ; and
it develops even with more vigor than in
the preceding cases. It usually rises as a
stout stem of several internodes lengthen-
ing almost simultaneously, or at least the
upper strongly developing long before the
lower have finished their growth; and
the latter are practically leafless, bearing
only small and scale-like and useless ru-
diments of leaves. This is correlated with
the peculiarity that the caulicle does not
lengthen in germination, or it lengthens
very slightly ; the cotyledons remain within
the coats of the seed; and if this were

40

buried beneath the surface of the ground, there it remains. The
abortion of the earliest leaves of the plumule is in correlation
with this Aypogeous (i. e. underground) situation of the cotyle-
dons throughout the germination. The slight elongation of the
caulicle serves merely to protrude its root-end from the coats of
the seed in a downward direction, and from this a strong root

usually is formed.

FIG. 39. Section of an acorn, filled by the embryo.
the same.

40. Advanced germination ot

20 MORPHOLOGY

33. In some Oaks, notably in our Live Oak (Quercus virens),
and less so in the Horsechestnut, the two cotyledons coalesce or
cohere by their contiguous faces.
In some of these cases of hypo-
geous germination, the short
ceaulicle and plumule are extri-
cated from the enclosing coats or
husk by the development of short
stalks (petioles, 157) to the fleshy
cotyledons ; as is seen in Fig. 42,
and in most germinating acorns.
These petioles are not visible in
the seed, but are the first develop-
ment in germination. ’

34. There are some curious
cases in which, while the caulicle
remains short and subterranean,
the cotyledons are raised out of
ground in germination by the
formation of far longer stalks
(petioles) than those of the
Horsechestnut. A singularly dis-
guised instance of this kind is seen in Megarrhiza, a genus of
Cucurbitaceous plants of California and Oregon, remarkable for
their huge root. The large seed has very thick and fleshy
cotyledons, and a very short and straight caulicle. In germi-
nation, the whole seed is elevated, seemingly in the manner of
the bean, upon a stout stem. One waits for a long time expect-
ing to see the cotyledons throw off the bursting husk and expand,
or else to put forth the plumule from between their bases. But
at length the plumule makes its appearance from an unexpected
place, coming separately out of the soil. Removing this, the
state of things represented in Fig. 43 is presented, — that of
the plumule seemingly originating from the base, instead of the
apex, of an elongated caulicle! But on examination of the cleft
from which this proceeds, by making a section of the stem above
(showing that it is hollow), and finally by separating the cotyle-
dons and gently tearing apart the two short stalks by which they
are united to their stem-like support, it is found that the latter may
be divided into two (as shown in Fig. 44), even down to the cleft
below. This explains the anomaly. The real caulicle has re-

FIG. 41. Section of a Horsechestnut or Buckeye seed. through the very thick
cotyledons and the incurved caulicle. 42. Seed in germination, showing the petioles
to the cotyledons, &c.

OF THE EMBRYO AND SEEDLING. 21

-mained short and subterranean, and is confluent with the upper
part of the thickening root: the seeming caulicle, which raised
the 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 germination it attains the
length of two or three inches: in age
it is readily separable into the two
leaf-stalks or petioles of which it is
composed: the plumule is thus seen
to be wholly normal, originating from
between the cotyledons. All the ex-
tensive growth so far, and until the
proper foliage-leaves of the continu-
ation of the plumule are developed
and begin their action, is from nutri-
tive material stored in the thickened
cotyledons, a considerable part of
which was transferred to the already
enlarging root, before a remaining
portion was used in building up the
strong plumule. The economy of this
elevation of cotyle-
dons which never’
open, and of the
lengthened distance
through which the
nutritive matter has
to be carried, is not
apparent. But it is
the family habit in
Cucurbitaceze to
bring up the cotyle-
dons that they may
develop as leaves
(as in the Pumpkin,
Fig. 47): here this
elevation is brought ut 43
about in a different way, but without securing the useful end.’

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

FIG. 43,44. Peculiar germination of Megarrhiza Californica; explained above.

a MORPHOLOGY

35. This same anomaly, as to the development of long stalks
to the cotyledons and their union into a stem-like body, occurs in
various species of Larkspur (notably in the Californian Delphin-
ium nudicaule) ; but in these the cotyledons develop into a pair
of efficient green leaves.

if

45
36. A similar elongation of petioles of the cotyledons, but
without any union, occurs in a species of Morning Glory of the
plains beyond the Mississippi (Ipomeea leptophylla) ; the leaf-
like cotyledons coming up on their long stalks separately from
the ground (Fig. 45); the developed plumule rising some
time afterward between them. Compare this with the ordinary
species (25, Fig. 15, 16, 25), and note that the difference is merely
that the caulicle in the common Morning Glory elongates and the
petioles of the cotyledons remain short.
37. In all instances thus far a single primary root so regularly
develops from the lower end of the axis of the embryo (variously
named radicle or caulicle), and forms such a direct downward

FIG. 45. Germination of Ipomca leptophylla; the caulicle not developing, the
plumule and the petioled cotyledons rise from underground. Dotted line marks the
level of the soil.

FIG. 46. Embryo of a Pumpkin, the cotyledons separated. 47. Same germinated ;
a cluster of roots from the base of caulicle.

OF THE EMBRYO AND SEEDLING. 23

prolongation of it, that it was called the descending axis; and
the body from which it originates was named the radicle, on the
supposition that it was itself the nascent root. But, as already
explained, the so-called radicle grows in the manner of stem (24),
and is morphologically that initial internode the node of which
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, Echinocystis, and the like,
the strong caulicle sends out directly from its root-end a cluster
of roots or rootlets, of equal strength; 7. e., it strikes root in
nearly the mainer that a cutting does. (Fig. 47.)
38. The Polycotyledonous Embryo is one having a whorl of
more than two seed-leaves. ‘The dicotyledonous embryo being
a whorl of the very simplest kind, that is, with the \
members reduced to two, the polycotyledonous |
may be regarded as a variation of it. In all but \ \ \
one group of plants it is simply a variation, of *
casual occurrence, or even a monstrosity, in which
three or rarely four cotyledons appear instead of
two. In Pines (Fig. 48, 49), however, and in
most but not all Coniferze, a whorl of from 3 to 10
cotyledons isshe normal structure, varying accord- ,
ing to the species, but of almost uniform number |
in oh. In germination these are brought out of»
the soil by the elongation of the caulicle, “and when
the husk of the seed is thrown off they expand K
into a circle of needle-shaped leaves. In the Pine * 49
tribe, all the subsequent leaves are alternate (spiral) in arrange-
ment, with some disguises. In the Cypress tribe, the cotyledons
are fewer (not more than four, and more commonly only two),
and the subsequent leaves also are in whorls of two to four;
z. e., are either opposite or verticillate. From the occasional
union at base of the cotyledons of a polycotyledonous embryo in
pairs or groups, and from a study of their early development,
Duchartre’ plausibly maintains that such cotyledons really consist
of a single pair, parted into divisions or lobes. The ordinary
interpretation, however, is equally tenable.

39. The Monocotyledonous Embryo, although theoretically the
simplest, is practically a more difficult study. It has a single
cotyledon (as the name denotes) ; also a single leaf to each node

1 Ann. Sci. Nat. ser. 3, x. 207. This view, which originated with Jussieu,
is adopted by Parlatore in DC. Prodr. xvi.

FIG. 48. Section of a seed of a Pine, with its embryo of several cotyledons. 49. Early
seedling Pine, with its stemlet, displaying its six seed-leaves.

24 MORPHOLOGY

of the plumule; that is, the leaves of the embryo are alternate.
But the caulicle is usually very short, and there is no external
mark by which its limits may be distin-
guished from the cotyledon, until germi-
nation has begun. For a type of it, the
embryo of some aquatic or marsh plants
may be taken, where it forms the whole
S A kernel of the seed (Fig. 50-53), and
50 51 52 53 the structure can be made out antecedent
to germination. It is understood by supposing that the cotyle-
don, which forms its principal bulk (the caulicle being only the
very short thickish base), is convolute around a short
plumule, and the margins conecreted, except a minute
longitudinal chink at base, out of which the growing
plumule protrudes in germination. The embryo of
Iris may be similar in structure, but no distinction
of parts is visible. It is very small in proportion to
the size of the seed, the kernel being mostly albu-
men, — a supply of food, from which the germinating
embryo draws the materials of its growth. When
this takes place, either the cotyledon or the whole
embryo lengthens, its lower part is pushed out of the
seed, a root forms at the free end of the excessively
short caulicle, and the plumule develops from the
other in a series of one-leaved nodes, the internodes
of which remain so short that the leaves continue
in close contact, the bases of the older successively
enclosing the inner and younger. (Fig. 55.) Here,
therefore, the cotyledon mainly remains in the seed,
and the seed remains underground (hypogzeous).
40. It is somewhat different in the Onion, which
has a similar embryo, except that it is longer, and
the cotyledon is curved in the albumen of the seed.
The first steps are the same as in Iris; but as soon
as a root is formed and embedded in the soil, the
cotyledon lengthens vastly more, into a long and
filiform green leaf, which, taking an erect position,

FIG. 50. Seed of Triglochin palustre; the rhaphe, leading to the strong chalaza at the
summit, turned towards the eye. 51. The embryo detached from the seed-coats, showing
the longitudinal chink at the base of the cotyledon; the short part below is the radicle.
52. Same, with the chink turned laterally, and half the cotyledon cut away, bringing to
view the plumule concealed within. 53. A cross-section through the plumule, more
magnified. Q

FIG. 54. Section of seed of Iris, enlarged, showing the small and apparently simple
embryo at the base of the albumen 55. Germinating seed and seedling of the same, of
natural size.

OF THE EMBRYO AND SKEDLING. 25

carries up the light seed far above the surface of the ground, the
tip only remaining in the albumen of the seed until that is ex-
hausted, when the tip perishes and the emptied husk falls away.
About this time the plumule shoots forth from one side of the
subterranean base of this cotyledonar leaf, in the form of a second
and similar filiform leaf, to be followed by a third, and so on.
The sheathing bases of these succeeding leaves become the coats
of the Onion-bulb. ‘The internodes remain undeveloped until the
plant is ready to blossom. Very similar is the germination of
a date-seed, except that the, 2 y} @, HLS
protruding cotyledon does_
not lengthen so much, nor/
does it elevate the heavy
seed. Instead of the seed
being carried up, the lower.’
end of the embryo, contain-
ing the plumule, is pushed down more or less into the loose
soil, from which in time the developing plumule emerges.

41. The embryo of Grasses, especially of those which yield
the cereal grains, is more complex, owing mainly to the great de-
velopment ‘of the plumule iT
and the manner in which «-~
its rudimentary leaves 3. |
successively enclose gr FAN
other. That of Maize or,
Indian Corn, one of the \W4
largest, is most convenient §© ° Gop s 4
for study. (Fig. 56-59.) The floury part of the seed, which
makes most of its bulk, is the albumen, largely composed of
starch. The embryo is exterior to this, applied to one of its
flat sides, and reaching from the thinner edge to or above the
middle in the common variety of corn here represented. The
form of the embryo is best shown, detached entire, in Fig. 58:
its structure appears in the sections. The outer part is the
cotyledon, which incompletely cnwraps the plumule: it adheres
closely to the albumen by the whole back, and remains un-
changed in germination: its function is to absorb nutritive

FIG. 56. Section, flatwise, of a grain of Indian Corn, dividing the albumen and the
embryo. 57. Similar section at right angles to the first. 58. A detached embryo:
corresponding parts of Fig. 57 and 58 indicated by dotted lines.

FIG. 59. Vertical section of Indian Corn across the thickness of the grain, dividing
the embryo through the centre and displaying its parts: ¢, cotyledon; p, plumule;
r, the radicle or caulicle.

FIG. 60. Similar section of grain of rice. 61. Same of an oat-grain; the parts
as in Fig. 59.

26 MORPHOLOGY

matter furnished by the albumen, and to transmit it to the.
growing plumule. ythe, giueues consists of a succession of
vudimentary leaves, sheathing
. and enclosing one another, on
the summit of a very short
-axis, which is mainly the
‘caulicle, otherwise called rad-
jicle. This is completely en-
closed by a basal portion of
the cotyledon and of the
-outermost leaf of the plu-
mie, which form a peculiar
‘sheath for it, named the
“Coleorhiza,* i. e. root-sheath :
_consequently the first root or
‘roots have to break through
‘this covering. As in the Oak
and Pea(32), the very first or
outermost leaves of the plu-
< mule develop imperfectly and not into
efficient foliage. The one in Fig. 62,
J which encloses the rest in the early
growth, is left behind as a mere sheath
to the base of the following and more
perfect leaves: it is the same as the
lowest in Fig. 63. The leaves are first
developed: the internodes lengthen later, and the lowest lengthen
very little. Not rarely the first root starts singly from the tip of
the caulicle (Fig. 62, just as in Fig. 55); but others of equal
strength follow from any part of the caulicle, and soon from
the nodes above; and no tap-root is ever formed.

42. A Pseudo-monocotyledonous embryo occasionally occurs ; .
that is, one of the dicotyledonous type, of which one cotyledon is
wanting through abortion. This occurs in Abronia, a genus
related to Mirabilis, and bearing an embryo very similar to that
represented in Fig. 17, 18, except that one cotyledon is absent.
The anomaly of an acotyledonous embryo occurs in Dodder, a
plant of the dicotyledonous type, but with both cotyledons

i\

2

1 This, the Coleorhize of Mirbel, should not be confounded (as by some it
has been) with the “root-cap,” or tissue which ordinary roots (whether
primary or secondary) break through in their development or carry on
their apex.

FIG. 62. Early germination of Indian Corn. . More advanced germination ot
same: roots produced from portion of stem above in ‘cotyledon as well as below.

OF THE ORGANS OF VEGETATION. 27

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

43. The dicotyledonous and the monocotyledonous character
of the embryo is correlated with profound differences in the whole
ulterior development, as revealed in the structure of the stem,
leaves, and flower ; which differences mark the two great divisions
of Phznogamous plants, viz. DicoryLepongs or DicoTyLEDONOUS
Priants, and MonocotyLepones or MonocoryLeponous Pants,
— names introduced into classification by Ray, and adopted by
A. L. Jussieu, in his Genera Plantarum.

CHAP TER - Ui:

MORPHOLOGY AND STRUCTURE OF THE ORGANS OF THE
PLANT IN VEGETATION.

Section I. Or tHe Roor.

44. The Root, which has been called the descending axis, is
that portion of the body of the plant which grows downward,
ordinarily fixing the vegetable to the soil, and absorbing from it
materials which the plant may elaborate into nourishment. As
already stated (24), the root grows in length by continuous
additions of new fabric to its lower extremity, elongating from
that part only or chiefly ; so that the tip of
a growing root always consists of the most
newly formed and active tissue. It normally
begins, in germination, at the root-end of the
caulicle, or so called radicle. But roots soon
proceed, or may proceed, from other parts of
the stem, when this is favorably situated for
their production. The root does not grow
from its naked apex, but from a stratum
immediately behind it: consequently its blunt
or obtusely conical advancing tip consists of older, firmer, and
in part effete tissue. The tip of all secondary roots and rootlets

_ FIG. 64. Magnified tip of root of a seedling Maple (such as in Fig. 9), sufficiently
enlarged to indicate the cellular structure: a. the portion where growth is taking
place; 5. the older and firmer tip.

28 MORPHOLOGY OF THE ROOT.

is similarly capped or protected.’ But the so-called root-cap is
seldom so distinct or separable as to deserve a particular name.

45. Nature of Growth, Cells. The development and growth of
the root, as of other organs, results from the development,
growth, and increase in number of certain minute parts, of which
the plant is built up. These component parts are so much alike,
at least in an early stage, and are so obviously formed all on
one type, that they take one common name, that of CELLs.
These are the histological elements of plants, 7. e. the units of
minute anatomical structure. While, in the morphology of the
plant’s obvious organs, analysis brings us to the phytomer (16)
as the individual element which by a kind of propagation
produces its like in a second phytomer, remaining however in
connection with the first, thus building up the general structure,
so, in an analogous way, each of the obvious parts — each stalk
or blade or rootlet —is microscopically determined to be com-
posed of these ultimate organic units, generally called cells.
The cell (cellula, by the French
conveniently termed cellule) is the
living vegetable unit, in the same
sense that the brick is the unit
of a brick edifice. To make this
analogy fairly complete, the
bricks should be imagined to
have a firm exterior or shell,
and a soft or at length hollow
interior, also to be living when
incorporated into the structure,
and finally to be produced in the
65 } forming structure by a kind of
«6 propagation. The production or
increase in number of these cells by development from previous
ones, and their successive increase in size up to maturity, are
what constitutes vegetable growth.2 The inspection through a

1 The notion that the tip of the root consists of delicate forming or
newly formed tissue, or bears some organ or structure of this nature (a
“ Spongiole”), has hardly yet been eliminated from the text-books and popular
writings. It had no proper foundation in fact.

In Lemna, and in some other aquatics, and also in some aerial roots, this
older tissue often separates into a real root-cap, free at base, like an inverted
calyptra.

2 This, as to the structure, is the subject of Histology; as to processes or
actions, the subject of Physiology ; both to be treated in a separate volume.

FIG, 65, 66. Portions of surface of Fig. 64, more magnified, clearly displaying the
superficial cellular structure and the long processes from some of the cells, called root-
hairs, whieh abound on the upper part of Fig. 64.

MORPHOLOGY OF THE ROOT. 29

simple microscope of a slender young root, and of thin slices
of it immersed in water, may serve to give a general though
crude idea of the vegetable cellular structure, sufiicient 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 predetermined order. The ultimate and
very slender branches are sometimes called root-fibrils ; but
these are only delicate ramifications of the root. Like any
other part of the plant, however, roots may produce hairs or
such like growths from the surface, which are wholly distinct
from branches. (3583.)

46. Root-hairs. Roots absorb water, &c., from the soil by
imbibition through the surface ; that is, through the walls of the
cells, which are in a certain sense permeable to fluids, more readily
when young and tender, less so when older and firmer. Roots,
therefore, absorb most by their fresh tips and adjacent parts ;
and these are continually renewed in growth and extended fur-
ther into the soil. As the active surface of a plant above ground
is enormously increased by the spread of foliage, so in a less
degree is the absorbing surface of young roots increased by the
production of root-hairs. (Fig. 64, upper part, and more magni-
fied in Fig. 65, 66.) These are attenuated outgrowths of some
part of the superficial cells into capillary tubes (only one from
each cell), closed at the tip, but the calibre at base continuous
with the cavity of the cell; into which, therefore, whatever is
imbibed through the thin wall may freely pass. These appear
(as Fig. 64 shows) at a certain distance behind the root-tip.
Further back the older or effete root-hairs die away as the cells
which bear them thicken into a firmer epidermis.

47. To the general statement that roots give birth to no other
organs, there is this abnormal, but by 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 power of originat-
ing them under certain circumstances, and some produce them
habitually. Thus Apple-trees and Poplars send up shoots from
the ground, especially when the superficial roots are wounded.
And the roots of Maclura or Osage Orange so readily originate
buds that the tree is commonly propagated by root-cuttings.

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

30 MORPHOLOGY OF THE ROOT.

latter are as normal as the primary root; that is, to stems
so situated that they can produce them. Most creeping plants
emit them freely, usually from the nodes; and so do most
branches, not too old, when bent to the ground and covered
with earth, thus securing the requisite moisture and darkness.
Separate pieces of young stems (cuttings) can commonly be
made to strike root. Upon this faculty of stems to originate
roots depends all propagation by division, by laying or layering,
by cuttings, &c. It is mainly annuals and common trees that
naturally depend on the primary root; and most of these can be
made to produce secondary roots. Even leaves and leaf-stalks
of some plants may be made to strike root and be used as
cuttings. (77.)

49. Duration. By differences in respect to this, cither the
root or the plant, as the case may be, is distinguished into
Annual, Biennial, or Perennial, according to whether life is contin-
ued for a single year or season, for two, or for a greater number.
The difference is not in all cases absolute or even weli marked.

00. Annuals are plants which, springing from the seed, flower
and seed the same year or season, and die at or before its close.
They produce fibrous roots, either directly from the embryo and
succeeding joints of stem (as in Grasses, Fig. 63), or from a
persistent primary or /ap-root, more or less thickened into a trunk’
or divided into branches. The products of vegetation in all such
herbs are not stored in subterrancan or other reservoirs, but are
expended directly in new vegetative growth, in the production
of blossom, and finally in the maturation of fruit and seed.
This completed, the exhausted and not at all replenished indi-
vidual perishes.

51. But some annuals may have their existence prolonged by
not allowing them to blossom or seed. Others, with prostrate
stem or branches, may from these produce secondary roots,
which, forming new connections with the soil, enable the newer
growth to survive when the older parts with the original root
have perished. And many herbs, naturally annuals, are continued
from year to year through such propagation from the branches,
used as layers or cuttings. Moreover, certain plants (such as
Ricinus or Castor-oil Plant), which 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 climates with comparatively warm and rainy
winter and rainless summer, many germinate in autumn, vegetate

MORPHOLOGY OF THE ROOT. 31

through the winter, flower and seed in spring, and perish in
early summer. These may be termed WryTER ANNUALS.

53. Biennials are plants which, springing from the seed and
vegetating in one season, live through the interruption of winter,
and ‘blossom, fructify, and perish in the next growing season :
their life being thus divided into two stages, the first of vegeta-
tion, the second of fructification. In typical biennials, nearly
the whole work of vegetation is accomplished in the first stage,
with the result of accumulation of a stock of nutritive matter,
to be expended in the second stage in the production of blossom
and seed. This accumulation is usually stored in the root or in
the base of a very short stem in connection with the root. The
root of a biennial accordingly enlarges and becomes fleshy, or
obese, as this matter accumulates. At the close of the growing
season, —‘the leaves perishing and the
stem having remained very short (with
undeveloped internodes), — the root,
crowned with the bud or buds, contains
the main result of the summer’s work,
as provision for the next year’s devel- \
opment and the completion of the
cycle. This development, being thus
amply provided for, is undertaken in
spring with great vigor ; blossom, fruit,
and seed are rapidly produced; and
the stock being consumed, but not at
all replenished, the cells of the great
root are now empty and effete, and
the individual perishes. The Beet,
Turnip, Parsnip, and Carrot are fa-
miliar examples of biennials, with the
store of nourishment in the root.?
The Kohl-rabi is a biennial with this deposit in the stem:
the Cabbage, partly in the stem, partly in the head of leaves.

1 In these the caulicle enlarges with the root, so that the upper and
bud-bearing end is stem.

Tap-roots of this kind are said, in descriptive botany, to be

Fusiform or Spindle-shaped, when broader in the middle and tapering
towards both ends, as in the common Radish (Fig. 67);

Conical, when tapering regularly from base to tip, as in carrots, &c. ;

Napiform, i. e. Turnip-shaped,when the thickened part is wider than high, &c.

Fascicled Roots are those which form in clusters ; these may be slender or
thickened. When much thickened, either irregularly or not of the above
shapes, they are said to be tuberous.

FIG. 67. Radish: a fusiform tap-root.

32 MORPHOLOGY OF THE ROOT.

54. But some plants, such as the Radish, which when they
spring from seed in autumn are true biennials, will when raised
in spring pass on directly to the flowering stage in summer, or
when sown after the warm season begins will often run through
their course as annuals. Then there are various biennials which
thicken the root very little and hold their leaves through the
winter. Between these and winter annuals no clear demarcation
can be drawn. As respects annual and biennial duration, the
terms may for the most part be applied indiscriminately to the
plant or to the root. We may say either that the plant is a
biennial, or that its root is biennial.

55. Perennials are plants which live and blossom or fructify
year after year. They may or they may not have perennial
roots. In trees and shrubs, also in
herbs with growth from year to year
from a_ strong tap-root, the root
is naturally perennial. But in most
perennials with only fibrous roots,
these are produced anew from time
to time or from year to year. Also,
while some such roots remain fibrous
and serve only for absorption, others
may thicken in the manner of the
ordinary biennial root and serve a
similar use, ¢. e. become reservoirs of
elaborated nourishment. The Dahlia
(Fig. 68) and the Peony afford good
examples of this. Sweet potato is -
another instance.t Most such roots
have only a biennial duration: they
are produced in one growing season; they yield their store to
form or aid the growth of the next. When perennials store up
nutritive matter underground, the deposit is more commonly
made in a subterranean portion of the stem, in tubers, corms,
bulbs, &c. (See 115-122.

56. The distinction between annuals and biennials is at times
so difficult, and the particular in which they agree so manifest,
—namely, that of blossoming only once, then dying, as it were
by exhaustion, —that it was proposed by DeCandolle to unite

1 It is only by the readiness of this root to produce adventitious buds,
especially from its upper part, that it has been mistaken for a tuber, such
as the common potato.

FIG. 68. Fascicled and tuberous or fusiform (secondary) roots of Dahlia: a,a. buds
on base of the stem

MORPHOLOGY OF THE ROOT. 33

the two under the common appellation of Monocarpric plants,
Plante monocarpice, taken in the sense of only once-fruiting
plants; and to designate perennials by the corresponding term
of Potycarpic, Plante polycarpice, literally many-fruited, taken
in the sense of many-times 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 species of
Lobelia should be ranked as annuals, biennials, or perennials.
The plants may blossom and seed toward the end of the season
in which they came from seed; or, germinated in autumn, the
small seedlings may survive the winter; but whenever fructified
the fibrous-rooted mother plant dies throughout; yet usually not
before it has established, and perhaps detached from the ba8e,
small offsets to blossom the next season; and so on. Then
Houseleeks (Sempervivum) and such-like fibrous-rooted succu-
lent plants multiply freely by offsets which are truly perennial
in the sense that they live and grow for a few or several years ;
but when at length a flowering stem is sent up producing blos-
som and seed, that plant dies as completely and in the same
manner as any biennial, only the generation of offsets surviving.
The same is true of the Century plant (Agave Americana,
wrongly denominated American Aloe), which yegetates in the
manner of the accumulating stage of a biennial, except that
this continues for several or very many years, while the flower-
ing stage, when it arrives, is precipitated and terminated in a
single season.

58. Although the stem usually sends forth roots only when
covered by or resting on the soil, which affords congenial dark-
ness and moisture, yet these are in some cases produced in the
open air. Roots may likewise subserve other and more special
uses than the absorption of crude or the storing of elaborated
nourishment.

59. Aerial Roots is a general name for those which are pro-
duced in the open air. One class of these may serve the office
of ordinary roots, by descending to the ground and becoming
established in the soil. This occurs, on a small scale, in the
stems of Indian Corn; the lower nodes emitting roots which
grow to the length of several inches before they reach the ground

1 These terms or some equivalents have a convenience in descriptive
botany. But those employed by DeCandolle are not happily chosen, as has
often been said. Mondtocous (bearing progeny once) and Polytocous (bearing
many times) would be more appropriate.

3

34 MORPHOLOGY OF THE ROO?Y.

into which they penetrate. More remarkable cases abound in
those tropical regions where the sultry air, saturated with moist-
ure for a large part of the year, favors the utmost luxuriance of
vegetation. In the Palm-like Pandanus or Screw-Pine? (Fig.
69), very strong roots, emitted in the open air from the trunk,
and soon reaching
the soil, give the
appearance of a tree
partially raised out
of the ground. The
famous Banyan-tree
of India (Fig. 71) is
a still more striking
illustration ; for the
aerial roots strike
from the horizontal
branches of the tree,
often at a_ great
height, at first swing-
ing free in the air,
but finally reach-
ing and establishing
themselves in the
ground, where they
increase in diameter
and form accessory
trunks, surrounding
the original bole and supporting the wide-spread canopy of
branches and foliage. Very similar is the economy of the Man-
grove (Fig. 70), which forms impenetrable thickets on low and
muddy sea-shores in the tropics throughout most parts of the
world, extending even to the coast of Florida and Louisiana.
Here aerial roots spring not only from the main trunk, as in
the Pandanus, but also from the branchlets, as in the Banyan.
Even the radicle of the embryo starts into growth, protrudes,
and attains considerable length while the fruit is still attached to
the branch.

59%. Aerial Rootlets for climbing are familiar in the Ivy of the
Old World (Hedera), Trumpet-Creeper (Tecoma radicans), and
our Poison Ivy (Rhus Toxicodendron) ; by the adhesion of

1 So named, not from any resemblance to a Pine-tree, but from a like-
ness of the foliage to that of a Pine-Apple.

FIG. 69. Pandanus, or Screw-Pine; and in the background, 70, a Mangrove-tree
(Rhizophora Mangle).

MORPHOLOGY OF THE ROOT. 30
which the stems, as they grow, ascend walls and the trunks of
trees with facility. In Rhus a superabundance of these rootlets
is produced, thickly covering all sides of the stem.

60. Epiphytes or Air-Plants also have roots which are through-
out life unconnected with the ground. Epiphytes, or Epiphytic
plants, as the name denotes, are such as grow upon other plants
without taking nourishment from them. Deriving this from the
air alone, they are called Air-plants. This name might be
extended to the same or other kinds of plants attaching them-
selves to bare walls, rocks, and the like, and unconnected with
the soil, though such would not technically be epiphytes. Very
many Lichens, Mosses, and other plants of the lower grade, and
not a few phznogamous plants, are in this case. The greater
part of the phenogamous Epiphytes pertain to two monocotyle-
donous orders, the Orchis family and that to which the Pine-
Apple belongs, viz. the Bromeliaceze. Their thread-like or
cord-like simple roots either adhere to the bark of the supporting
tree, securing the plant in its position, or some hang loose in the
air. Of these, Orchids, 7. e. plants of the Orchis family, are the
most showy and numerous, and of the greatest variety of forms,
especially of their blossoms, which are often bizarre and fantas-
tic. They belong, naturally, to climates which are both warm
and humid: they are highly prized in hot-house cultivation ;
and, along with the hardy and terrestrial portion of the order,
they are peculiarly interesting to the botanist on account of the
singular and exquisite adaptation of their flowers in relation
to insects which visit them. In some the blossoms curiously

FIG. 71. The Banyan-tree, or Indian Fig (Ficus Indica).

36 MORPHOLOGY OF THE ROOT.

resemble butterflies or other insects; as, for example, Oncidium
Papilio, Fig. 72. Epiphytic orchids are indigenous to the United
States only from Georgia to Texas, and only in humble forms,
in company with species of Tillandsia, representing Bromeliace~
ous epiphytes. The commonest of the latter tribe, and of most
northern range, is the T. usneoides, the so-called Long Moss,
which, pendent in long and tangled gray clusters or festoons from
the branches of the Live-Oak or Long-leaved Pine, gives such a
peculiar and sombre aspect to the forests of the warmer portions
of our Southern States. =

61. Parasitic Plants have the peculiarity that their roots, or
what answer to roots, not only fix themselves to other plants,
but draw therefrom their nourishment, at least in part. Among
cryptogamous plants very many Fungi are parasitic upon or
within living plants or animals. But only pheenogamous para-
sites are here under consideration. These may be divided into
two classes ; those with and those without green foliage.

62. Green Parasites may be either wholly or partially parasitic ;
that is, they may draw all their support from a foster plant, or

FIG. 72. Oncidium Papilio, and, 73, Comparettia rosea; two showy epiphytes of the
Orchis family; showing the mode in which these Air-plants grow.

MORPHOLOGY OF THE ROOT. 37

they may be likewise rooted in the soil, and receive from it
materials 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 examples of complete green
parasitic plants. ° Seeds dropped by birds on the boughs of trees
germinate there ; the root-end of the caulicle points thither instead
of towards the earth; the root, or what would be such, pene-
trates the bark and in-
corporates itself with the
sap-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 is, both
by what the foster tree
has assimilated and £7
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. Partially parasitic plants (mostly green) may be either woody
and arborescent or herbaceous. The species of Clusia in tropical

\\

PY
‘i
¥

iss

FIG. 74. Native epiphytes of Georgia, &c.: the erect one at the right an Orchid,
Epidendrum conopseum; the hanging one Tillandsia usneoides, called Long Moss.

FIG. 75. Roots of Gerardia flava: some of the rootlets attaching themselves para-
sitically to the root of a Blueberry. (From @ drawing by Mr. J. Stauffer.)

3 MORPHOLOGY OF THE ROOT.

America (called Cursed Fig) are examples of the former. They
form trees, send down aerial roots in the manner of the Banyan ;
but, while some roots seek the ground, some may attach them-
selves to other trees parasitically, and draw from them a portion
of their support. The parasitism of certain herbaceous plants
with green foliage is clandestine, the connection being under-
ground and therefore long unsuspected. This occurs in species
of Gerardia (at least of the section
Dasystoma) and other plants of the
same family, the uncultivability of
Which is thereby explained. Also
@) in Comandra and in their relatives
‘the Thesiums of the Old World,
belonging to a aie order (the Santalaceze) which has much
affinity aie the entirely parasitic order (Loranthacex) to which
the Mistleto belongs.

64. Pale or Colored Parasites, such as Beech-Drops, Pine-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
from the sap already elaborated.
They have accordingly no occasion
for digestive organs of their own,
2.e. for green foliage. The Dodder
(Fig. 77) is a common plant of
this kind which is parasitic above
ground. Its seeds germinate in
the earth, but form no proper root:
when the slender twining stem
: reaches the surrounding herbage,

7 78 7 it forms suckers, which attach
themselves firmly to the surface of the supporting plant,
penetrate its epidermis, and feed upon its juices; while the
original root and base of the stem perish, and the plant has
no longer any connection with the soil. Thus stealing its nour-

FIG. 76. Section of one of the attached rootlets of Gerardia, showing the union. +

FIG. 77. The common Dodder of the Northern States (Cuscuta Gronoyii), of the
natural size, parasitic upon the stem of an herb: the uncoiled portion at the lower end
shows the mode of its attachment 78. The coiled embryo taken from the seed, con-
sisting of naked caulicle and plumule; moderately magnified. 79. The same in germi-
nation, elongating into a thread-like leafless stem.

2

MORPHOLOGY OF THE ROOT. 39

ishment ready prepared, it requires no proper digestive organs
of its own, and, consequently, does not produce leaves. This
economy is foreshadowed in the embryo 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 flax-fields, having 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 indifferently 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 Pilostyles (para-
sitic flowers on the shoots of Leguminous plants) in Tropical
America, one species of which was discovered by Dr. Thurber

near the southern borders of New Mexico. Its flowers are
small, only about a quarter of an inch in diameter. The most
wonderful plant of this kind is that vegetable Titan, the Raf-
flesia Arnoldi of Sumatra (Fig. 80) which grows upon the stem
of a kind of a Cissus or Vitis. It is a parasitic flower, measuring
nine feet in circumference, and weighing fifteen pounds! Its
color is light orange, mottled with yellowish-white.

1 Monotropa or Indian Pipe (and perhaps some related plants), although
probably parasitic on living roots in early growth, appears to live afterwards
in the manner of the larger Fungi, upon leaf-mould and decaying herbage.
Its mode of life should be investigated.

FIG. 80. Rafilesia Arnoldi; an expanded flower, and a bud, directly parasitic on
the stem of a vine: reduced to the scale of half an inch to a foot.

40 MORPHOLOGY OF BUDS.

Section II. Or Bups.

65. Buds are the germs of stems: they are axes with their
appendages in an early state. Lrar-sups (GEMM) are those
devoted to vegetation, and the parts, or some of them, develop
as leaves. Mixep subs contain both foliage and flower or
flowers. FLOWER-BUDS (ALABASTRA) are unexpanded blossoms.
These are considered in another chapter.

66. The conspicuous portion of an ordinary bud, or that which
first develops, usually consists of leaves, or scales the homologues
of leaves; the axis itself being very short and undeveloped. If
this remains comparatively short, the leaves as developed are
crowded in a rosette, as in a Houseleek (Fig. 91°), a Barberry
and the Larch: when the internodes lengthen, the leaves are
interspaced upon the axis.

67. The cotyledons and plumule of the embryo are, morpho-
logically, the first bud, on the summit of the initial stem, the
caulicle. This in germination and subsequent growth develops
into a leafy stem, in the manner already described. Normally
this stem has the capacity of growing on in this way from the
apex or growing point, which is always potentially a bud, the
apical or terminal bud (15). Sometimes it is merely potential,
and there is no external structure visible until the new growth
begins, or the bud is said to be Jatent.

68. But commonly, in plants that live from year to year, growth
is divided into seasons or stages, with intervals of repose. In
such cases, especially in trees and shrubs, instead of a continuous
succession of foliage, the period of interruption is apt to be
marked by the production of scales (Bud-scales, Perule, ete.) or
dry teguments, which serve to protect the tender rudiments or
growing point within during the season of rest. This being the
winter-season in cold climates, Linnzeus gave to such bud-coyer-
ings the common name of Hisernacutum. From the usually
squamose (scale-like) character of this covering, such buds take
the name of

-69. Sealy Buds. Large and strong ones of this kind, such as
those of Horsechestnut, Magnolia, Hickory, Lilac, &c., may be
taken as the type of bud. The scales serve to protect the ten-
der parts within against injury from moisture and from sudden
changes in temperature during the dormant or earliest growing
state. To ward off moisture more effectually, they are sometimes
coated with a waxy, resinous, or balsamic exudation, as is con-
spicuous on the scales of the Horsechestnut, Balsam-Poplar or
Balm of Gilead, and Balsam-Fir. To guard against sudden

MORPHOLOGY OF BUDS. 41

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

70. Nature of Bud-seales. That they answer to leaves is made
manifest by « consideration of their
situation and arrangement, which are
the same as ofthe proper leaves of
the species ; and by the gradual transi-
tions from the former to the latter in
many plants. In the 7urions, or sub-
terranean budding shoots of numerous
perennial herbs, and in the unfolding
buds of the Lilac and Sweet Buckeye
(4Esculus parviflora), every gradation
may be traced between bud-scales and
foliage, showing that no line of distinc-
tion can be drawn between them, but
that the two are essentially of the same
nature, are different modifications of
the same organ. In the Lilac they
may be regarded as the blade of the
leaf, modified and depauperate ; in the
Buckeye (Fig: 253), and therefore in
Horsechestnut, as the base of leaf-stalks;
in Magnolia (Fig. 81, 82), in the
Tulip-tree, and in the Beech, they are
evidently stipules. They must therefore
be referred to in the section on the
morphology of leaves. (227.)

71. Naked Buds, &c., 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 leaves of the season in Viburnum
nudum, V. lantanoides (Hobblebush),
and the like, remain in a nascent state
over winter without covering, and ex-
pand into the first foliage in the spring.
Yet V.Opulus (Snowball, &c.), another species of the same
genus and inhabiting the same region, has well-formed scaly

FIG. 81. Branch of Magnolia Umbrella, of the natural size, crowned with the
terminal bud; and below exhibiting the large rounded lJeaf-scars, as well as the rings
or annular sears left by the fall of the bud-scales of the previous season. 82. A detached
scale from a similar bud; its thickened axis is the base of a leaf-stalk ; the membranous
sides consist of the pair of stipules united with it.

42

leaf-buds.

MORPHOLOGY OF BUDS.

In other hardy shrubs and trees, the buds, equally

or almost destitute of scales,
are minute, hidden in or un-
der the bark, or otherwise
inconspicuous until vernal
growth commences. Phila-
delphus and Taxodium are of
this kind.

72. Subpetiolar Buds. Some
leaf-buds are singularly cov-
ered in their early state and
through the summer, as in the
Locust (Robinia), Honey-Lo-
cust Fig. 96 (where they re-
main very undeveloped), in
Yellow Wood (Cladrastis). and
more conspicuously in the
Plane-tree (Platanus, Fig. 87):
here they are all formed un-
der the base of the protecting
leaf-stalk, which in Plane-tree
forms a sheath or inverted cup,
very like a candle-extinguisher,

fitted to and concealing the conical bud until autumn, when by
the fall of the leaves these buds are exposed.

FIG. 83.

Diagram of vértical section of a strong bud, such as of Horsechestnut.

84. The axis of the same developing, the elongation beginning with the lowest inter-

node, soon followed by the others in succession. 85. A year’s growth of Horsechestnut,

crowned with a terminal bud: a, scars left by the bud-scales of the previous year: ~
6, scars left by the fallen leaf-stalks: ec, axillary buds.

FIG. 86.
FIG. 87.

Branch and buds (all axillary) of the Lilac.
Leaf-bud under the petiole of the Plane-tree.

MORPHOLOGY OF BUDS. 43

73. Fleshy Buds. ulbs are peculiar buds of certain herba-
ceous plants, with fleshy scales, and often of a more permanent
character. Their nature and economy may most conveniently
be illustrated under subsequent sections. Usually bulbs are
subterranean or partly so. But small bulbs ( Bulblets, 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
ground, produce roots, and grow as independent plants ; and
their fleshy 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 bulblets or buds
establishing themselves as progeny. In several species of
Allium (Onions and Leeks), such bulblets 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 grow independently. 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 root into the soil
and becoming a separate plant. The horticultural advantage of
bud-propagation is, that the offsets or new individuals share
in all the peculiarities of the parent as completely as if still
branches of that tree. In propagation by seed, the special
peculiarities or excellencies of individuals or varieties may not,
and im some measure probably will not, be reproduced.

75. Normal or Regular Buds, as to position, are either terminal
or axillary, 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 upper angle
formed with the stem by) any well-developed leaf. In these
positions, buds are so usual, or so capable of appearing, that
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

44 MORPHOLOGY OF BUDS.

surmounts: axillary and any other lateral leaf-buds are to be-
come branches. But even of buds which actually appear a large
proportion do not grow. When a terminal bud is formed (as
Fig. 81, 85, 91), this is commonly the strongest, or among the
stronger. But in many cases it habitually or commonly fails to
appear. In the Elm (with leaves and therefore buds alternate),
the bud axillary to the last leaf of the season takes its place.
In the common Lilac, a pair of buds, which were in the axils of
the uppermost of the (opposite) leaves, seem to replace the
terminal bud, which seldom develops. (Fig. 86.) When all the
regular buds make their appearance, and the leaves are opposite,
the stem will be crowned with the terminal bud, having an axil-
lary bud on each side of it. (Fig. 88.)

76. Aecessory Buds. These are, as it were, multiplications of
the regular axillary bud, giving rise to two, three, or more, instead

of one; in some cases situated one above an-
other (sperposed ), in others placed side by
side (collateral). In the latter case, which
occurs occasionally in the Hawthorn, in cer-
tain Willows, in the Maples (Fig. 88), &e.,
the axillary bud seems to divide into three,
or itself to give rise to a lateral bud on each
side. On some shoots of the Tartarean
Honeysuckle (Fig. 90) from three to six buds
appear in each axil, one above another, the
lower being successively the stronger and
earlier produced; and the one immediately
in the axil, therefore, grows in preference:
occasionally two or more of them grow, and
superposed accessory branches result. It is
much the same in Aristolochia Sipho, except
that the uppermost bud is there strongest.

FIG. 88. Branch of Red Maple, at the middle bearing triple axillary buds, placed
side by side.

FIG. 89. Piece of a branch of the Butternut, with accessory buds placed one above
another: a, the leaf scar: 6, proper axillary bud: ec, d, accessory buds.

FIG. 90. Part of a branch of Tartarean Honeysuckle, with crowded accessory buds
superposed in the axil of each leaf.

MORPHOLOGY OF STEMS. 45

So it is in the Butternut (Fig. 89), where the true 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 ewtra-axillary branch.

77. Adventitious 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, especially when wounded, and in some
plants (such as Blackberries) so freely that gardeners propagate
them by 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 produced on the margins 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 II. Or tHe Srem.
§ 1. GENERAL CHARACTERISTICS AND GROWTH.

78. The Stem is the ascending axis, or that portion of the
trunk which in the embryo grows in an opposite direction from
the root, seeking the light, and exposing itself as much as pos-
sible to the air. All phaznogamous plants possess stems. In
those which, in botanical descriptions, are said to be acaulescent,
or stem/ess, 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 uniformly retain
it; but sometimes trails along the surface of the ground, or
burrows beneath it, sending up branches, flower-stalks, or leaves
into the air. The common idea, that all the subterranean portion
of a plant belongs to the root. is incorrect. Equally incorrect 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 Wolffia can the stem be said to be wanting.

46 MORPHOLOGY OF STEMS.

79. While the root normally gives birth to no other organs,
but itself performs those functions which pertain to the relations
of the vegetable with the soil,— binding it to the earth and
absorbing nourishing materials from it, — the aerial functions of
vegetation are chiefly carried on, not so much by the stem it-
self as by a distinct set of organs which it bears, namely, the
leaves. Hence, the production of leaves is one of the charac-
teristics of the stem. These are produced only at certain definite
and symmetrically arranged points, called nodes. (13, 23.)

80. Development and Structure. Ina bud or undeveloped stem,
the nodes are in contact or close proximity. In the develop-
ment, growth in length takes place in such manner as to carry
these apart more or less, according to the degree of elongation,
that is, the internodes (13) elongate. The order of development
is from below upward, the lowest internode 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. Unlike the root, in which the
elongation of formed parts is very soon finished and therefore
only the tip is perceptibly growing, internodes go on growing
throughout, and several formed internodes may be growing
simultaneously, thus producing elongation throughout a consid-
erable extent of stem and with considerable rapidity. But each
internode grows independently. Some parts of an internode
may lengthen faster or continue in growth longer than others ;
this is usually the upper portion, at least in long internodes and
when every part is equally exposed to light.

81. The deveiopment of a stem from a bud is wholly like that
from the embryo, and has already been described in Chap. II.
It exhibits similar variations as to rapidity and vigor, dependent
upon the constitution of the bud, — which, like the plumule in
the seed or seedling, may be either latent or much developed
before growth begins, —also upon the amount of nourishment
provided. Strong buds commonly have their parts, or some of
them, ready formed in miniature, and a store of elaborated 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 the preceding
summer (as in Magnolia, Fig. 81, Horsechestnut, Fig. 85, and
Hickory, Fig. 91) often exhibit the whole plan and amount of
the next year’s growth; the nodes, the leaves they bear, and
sometimes the blossoms being already formed, and only requiring
the elongation of the internodes for their full expansion. As

CHARACTERISTICS AND GROWTH. 47

the bud is well supplied with nourishment in spring by the stem
on which it rests, its axis elongates rapidly ; and although the
growth commences with the lowest internode, yet
the second, third, and fourth internodes may
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), they
are the uppermost axillary, which take the place of
a suppressed or abortive terminal bud.

82. Such woody stems, developed from a strong
bud, and terminated at the close of the season’s
growth by a similar bud, may be continued from
year to year in an unbroken series. 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 they 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-scars, or marks on the bark left by the separation
of the leaf-stalk, which are for a long time conspicuous on the
shoots of the Horsechestnut (Fig. 85 6), the Magnolia (Fig. 81),
and Hickory, Fig. 91.

83. Ramification. Brancues (14-16) are secondary stems
developed from a primary one, or tertiary ones from these, and
soon. Ultimate or small ramifications of latest order are some-
times called BrancuLets. The terminal bud continues the stem
or axis which bears it. Lateral buds give rise to branches.’
As the normal lateral buds are axillary (75), so are normal
branches. The symmetry 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 phenogamous plants that it may here be left
out of view.

In phenogamous 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 leaves and their modifications.

FIG. 91. End of a Hickory branch (Carya alba), with a strong terminal and smaller
axillary buds.

48 MORPHOLOGY OF STEMS.

branches wiil be alternate; when the leaves are opposite, and
the buds develop regularly, the branches will be opposite, &e.
This holds in fact sufficiently to determine and exemplify the
plan of ramification ; but, if entirely carried out, there would be
as many branches as leaves. ‘This could rarely if ever be, even
in primary ramification.

84. Non-development of Buds. Someof the buds are latent or
merely potential, that is, do not make their appearance : of those
which do appear only a part actually grow into branches ; and
of these some are apt to perish at an early stage. In our trees,
most of the lateral buds generally remain dormant for the first
season: they appear in the axils of the leaves early in summer,
but do not grow into branches until the following spring; and
even then only a part of them grow. Sometimes the failure
occurs without appreciable order; but it often is nearly uniform
in each species. Thus, when the leaves are opposite, there are
usually three buds at the apex of a branch ; namely, the terminal,
and one in the axil of each leaf; but it seldom happens that all
three develop at the same time. Sometimes the terminal bud
continues the branch, the two lateral generally remaining latent,
as in the Horsechestnut (Fig. 85) ; sometimes the terminal one
fails, and the lateral ones grow, when the stem annually becomes
two-forked, as in the Lilac, Fig. 86. The undeveloped buds
do not necessarily perish, but are ready to be called into action
in case the others are checked. When the stronger buds are
destroyed, some that would else remain dormant develop in their
stead, incited by the abundance of nourishment, which the for-
mer would have monopolized. In this manner our trees are soon
reclothed with verdure, after their tender foliage and branches
have been killed by a late vernal frost, or consumed by insects.
And buds which have remained latent for several years occasion-
ally shoot forth into branches from the sides of old stems,
especially in certain trees.

85. Most branches springing from old trunks, however, as in
Willows and Poplars, especially when wounded or pollarded,
originate from adventitious buds (77), which occur without
order. So also when accessory buds (76) develop into branches,
normal symmetry is more or less disturbed, as by contiguous
shoots standing directly over each other in Tartarean Honey-
suckle, or by a branch far out of the axil in Walnuts (Fig. 89)
and Honey-Locust, Fig. 96.

86. Excurrent and Deliquescent Stems. Sometimes the primary
axis is prolonged without interruption, even through the whole
life of a tree (unless accidentally destroyed), by the continued

CHARACTERISTICS AND GROWTH. 49

evolution of a terminal bud, or by some upper strong bud which
equally 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 del/quescent, as in the White Elm and most of
our deciduous-leaved trees. The first naturally gives rise to coni-
cal or spire-shaped 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 strikingly 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 of the trunk.

87. Definite and Indefinite Annual Growth of Branches. In
many of our trees and shrubs, especially those with scaly buds,
the whole year’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 completed long before the end
of summer ; when the shoot is crowned with a vigorous terminal
bud, as in the Horsechestnut (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 directly, 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 branches
grow onward indefinitely, until arrested by the cold of autumn:
the buds at or near their summit are consequently young and
nnmatured, or at least the lower and older axillary buds are
more vigorous, and alone develop into branches the next spring ;
the later-formed upper portion most commonly perishing from
the apex downward for a certain length in the winter. The
Rose and Raspberry, and among trees the Sumac and Honey-
Locust, are good illustrations of this sort; and so are most
perennial herbs, their stems dying down to or beneath 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
vegetation.

88. Many of the details and applications of ramification, of
most importance in morphology and descriptive botany, relate

50 MORPHOLOGY OF STEMS.

to anthotaxy or inflorescence (Chap. V.), which has its own
terminology. But some of its terms may be conveniently
employed in the description of ramification unconnected with

flowering.

§ 2. Forms or STEMS AND BRANCHES.

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

90. Herbs are plants in which the stem does not become
woody and persistent, but dies annually or after flowering, down
to the ground at least. The difference between annual, biennial,
and perennial herbs has already been pointed out in the chapter
on the root (50-57), and the gradations between them indicated.
Herbs pass into shrubs and shrubs into trees through every gra-
dation. The following definitions are therefore only general : —

91. Undershrubs, or Suffruticose plants, are woody plants of
humble stature, their stems rising little above the surface. If
less decidedly woody, they are termed Suffrutescent.

92. Shrubs are woody plants, with stems branched from or
near the ground, and less than five times the height of a man.
A shrub which approaches a tree in size, or imitates it in aspect,
is said to be Arborescent.

93. Trees are woody 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 from the ground to the
height of thirty feet or more, may still be called shrubs.

94. The erect position, elevation above the soil, and self-sup-
port, are normal conditions of the stem, but are far from universal.
And certain kinds of stem or branches are sufliciently peculiar
to have received substantive names: other equally peculiar forms
have no special names. There are, moreover, certain organs
(such as spines and tendrils) which are commonly homologous
(12) with stems, but not always. ‘Two kinds of erect stems
have special names in descriptive botany.

95. Culm is a name applied to the peculiar closed-jointed stem
of Grasses and Sedges, whether herbaceous, as in most Grasses,
or woody or arborescent, as in the Bamboo.

96. Caudex is the name technically applied to the trunk of
Palms (Fig. 126), Tree-Ferns, and the like, consisting of a
commonly simple column, the surface beset with scales, — the

SPECIAL FORMS. oh

bases of former leaf-stalks, — or marked by scars, left by their
fall. This name was used by botanists anterior to Linnzeus for
any tree-trunk, but is now used for the peculiar stems above-
mentioned ; also for the persistent base of a stem, otherwise
annual, which throws up fresh herbaceous stems or stalks from
year to year. Such short and enduring stems, being usually
near the ground. or under it, were commonly 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 Seape 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. (265.) Scapes
usually spring from some one of the subterranean forms of stem.

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

99. Seandent or Climbing Stems are those which rise by
attaching themselves to some extraneous support. ‘This is
effected in various ways; in some by the action of the stem
itself, in others by that of organs which it bears.?

100. Voluble or Twining Stems, or Twiners, are those which
ascend by coiling round a support, which must accordingly be
comparatively slender, or at least not too large. Some ascend
by coiling ‘‘ with the sun” (that is, from right to left of the
observer viewing the coil from the outside”), as the Hop; more,

1 See Darwin, The Movements and Habits of Climbing Plants, London
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 are in popular
language called Vines (e. g. Hop-Vine, Grape-Vine, Squash-Vine, &c.), a
name which properly belongs to Vitis only.

2 Dertrorse and Sinistrorse, i. e. to the right or to the left, are 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. 891) suggested Eutropic for the former, Antitropic 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. Linneus supposed the observer to view the

52 MORPHOLOGY OF STEMS.

by coiling in the opposite direction, as the Bean (Phaseolus),
the woody Aristolochia Sipho, the Morning Glory (Fig. 91*) and

other Convolyu-
lace. The Dod-
der, a leafless par-
asitic plant of the
latter family, not only gains suppoit by coiling
on the stems of other plants, but by attachment,
through the development of sucker-like discs,
along the whole contiguous surface. (Fig. 77.)
The various actions through which plants climb,
and the attendant phenomena, are physiological,
and will be treated in the second part of this
Text Book. The most complete and satisfac-
tory discussion of the subject, of a readable sort,
is that of Darwin’s volume, referred to in a

ola preceding note.

101. Leaf-Climbers 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,
Tropzolum, and Solanum jasminoides (Fig. 255) ; in some by
the incurvation of leaf-blades or portions of them, as in Adlu-
mia; or by an extension of the midrib into a hook or short ten-
dril, as in Gloriosa; or by the transformation of some of the
blades of a compound leaf into hooks or tendrils, as in Cobzea
and the Pea.

102. Tendril-Climbers (Fig. 92-95) are those in which the
prehension is by a tendril, a slender filiform body, either simple
or branched, specially adapted to the purpose, and capable of
coiling, either to secure a hold, or to draw the stem up to the

coil or circle from the inside; Mohl, Palm, Braun, and the DeCandolles
adopt this, and the latter insist on it. Such authority should be decisive,
if common usage and popular sense went along with it. But some of the
botanists following Linneus adopted the reverse view; and to the present
writer, as to Bentham and Hooker, Darwin, Eichler, and in part G. Henslow,
it was so natural to view the coil from the outside that we without concert
adopted this position and mode of expression. A right-hand coil, or one
turning to the right, with us, is one the turns of which pass from the left to
right of a bystander who confronts the coil. It is in this sense that a com-
mon screw is called a right-handed screw, and that the right bank of a river
is that to the right of the person who follows the course of the stream. So
natural is this, that even on a map or plate, which has face and back, and
therefore a right and left of its own, the figures occupying its right or left
portions are understood to be those which are toward the right or the left
hand of the observer who stands before it.

FIG. 91%. Dextrorsely twining stem of Morning Glory, Ipomca purpurea.

vet Te
eo

SPECIAL FORMS. 53

support. In certain tendrils the attachment to the support is by
a sucker-like disc at the apex, as in the Virginia Creeper or
Ampelopsis, Fig. 94.

103. Root-Climbers are those in which the stems produce aerial
rootlets (597), which fix themselves to a supporting surface
along which the stem creeps or ascends. In this way Trumpet
Creeper (Tecoma radicans), Ivy, and Poison Ivy (Rhus Toxi-
codendron) climb extensively.

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

Ascending or Assurgent, when they rise obliquely upward ;

Reclining, when from an ascending or erect base the upper
part recurves and trails ;

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

Procumbent or Prostrate, when lying at length upon the ground ;

Repent or Creeping, when growing prostrate on the ground
and rooting as they grow. Also applied to similar stems grow-
ing under, as well as upon the surface of the soil, as in Couch-
Grass and Mint, Fig. 99.

105. A Sucker (Sureulus) is an ascending stem rising from a
subterranean creeping base. The Rose and Raspberry multiply
freely by suckers. Such plants are easiest to propagate ‘+ by
division.”

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

. 107. An Offset is a short stolon or a short sucker. Houseleek
(Fig. 91") offers a familiar
example. By offsets, some
herbs, otherwise annuals, are
continued from year to year in
a vegetative progeny (Lobelia
cardinalis, &c.), and peren-
nials may thus establish colo-
nies around a parent individual.

108. A Runner (Flagellum)
is a filiform or very slender
stolon, naked and tendril-like except at tip, where it roots,
develops a bud, and so a new plant. The Strawberry furnishes
the most familiar example.

FIG. 914. Houseleek (Sempervivum tectorum) with offsets.

54

MORPHOLOGY OF STEMS.

109. The two following are organs which may be of axial
nature, or may not. This may ordinarily be determined by posi-

tion. Any direct continuation of
stem or branch must be of axial
nature, that is, of the nature of stem ;
and the same is true of whatever
primarily develops in the axil of a
leaf. Conversely, whatever subtends
a lateral axis or branch may be taken
for a leaf or foliar production, being
in the place of such.

“410. A Tendril, a thread-shaped

~ and leafless body, capable of coiling

spirally, and used for climbing (102),
is homologous with stem in Grape-
vines (Fig. 92) ; for the uppermost
tendril is seen to be a direct continu-
ation of the stem. ‘The small bud
which appears in the axil of the
uppermost leaf will in its growth
produce another internode and leaf,
or some species more than one,
but will terminate in a_ similar

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

FIG. 92. End of a shoot of the Grape-vine, with young tendrils: a sympodial

stem. (See note.)

FIG. 93. A portion of a stem of Ampelopsis quinquefolia, or Virginia Creeper, with

a leaf and a tendril.

FIG. 94. Ends of the latter, enlarged, showing the expanded tips or dises by which

they cling.

t
dt

-
-
~

SPECIAL FORMS.

the figure.’ The tendrils of Virginia Creeper (Fig. 93) are of the
same nature and position. But, instead of laying hold by a coiling
of the tip, when it has reached any solid surface, such as a wall
or tree-trunk, the tip expands into an adhesive disc, which forms
a secure attachment. (Fig. 94.) Ina related plant, Vitis (Cissus)
tricuspidata of Japan, these disks terminate the branches of very
short tendrils: consequently the shoots as they grow are at
once applied closely and secured firmly to the surface of the sup-
port, — an admirable adaptation for climbing walls and trunks.
N

111. The simple tendril of a Passion-
flower, being in the axil of a leaf (that
isin the position of a branch), is also of
axial nature: it is a leafiess and simple
branch, composed of one long and slen-
der internode, devoted to the purpose of
climbing. Fig. 95 shows in all stages the
admirably active tendrils of Passiflora
sicyoides. This is a Mexican species,
remarkable for the rapidity and freedom
with which the tendrils move. The lowest
tendril in the figure is attached and
coiled: the next is free and coiled in
one helix: the third is outstretched
and seeking a support. For tendrils
‘= =owhich are not homologous with stems,

95 "see Sect. IV. 228.

112. A Spine or Thorn (Fig. 96, 97) is usually a branch or
the termination of a stem or branch, indurated, leafless, and
attenuated to a pomt. The nature of spines is manifest in the
Hawthorn (Fig. 97), not only by their position in the axil of a
leaf, but often by producing imperfect leaves and buds. And
in the Sloe, Pear, &c., many of the stinted branches become
spinose or spinescent at the apex. tapering off gradually into a
rigid and leafless point, thus exhibiting every gradation between a
spine and an ordinary branch. These spinose branches are less

1 This forms what is called a Sympodium or Sympodial stem, which is mor-
phologically made up of a series of superposed branches. (See Chapter V.
281,282.) In contradistinction, a stem formed by the continued development
of a terminal bud is Monopodial or a Monopodium. Fig. 95 is an example.

FIG. 95. Leafy shoot of Passiflora sicyoides, of Mexico, with fixed and coiled, free
and full grown, and forming tendrils.

56 MORPHOLOGY OF STEMS.

liable to appear on the cultivated tree, when duly 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
(76), one on each side, ap-
pear, and grow the next sea-
son into ordinary 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.

115. 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
superiicial outgrowths from
the bark, of? a different nature
(385), and of small morpho-
logical signification.

114. Subterranean Stems are hardly less diverse than the
aerial. They are classed as Ruizomes, Tusers, Corms, and
Buss, the forms passing one into another by gradations.

115. Rhizoma (/hizome, or in English Roorsrocxk) 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 coy-
ered by the soil, and usually produces from its apex some kind
of aerial stem, either leafy or as a flower-stalk (scape, 97),
which rises into the air and sight. . Before morphology was
understood, rootstocks were called creeping roots, scaly roots, &e.
Some are slender, such as those of Mints (Fig. 99), of most
Sedges (Fig. 98), and of Couch-Grass. Their cauline nature is
evident from their structure and appearance; their nodes and
internodes are well marked, the former bearing leaves reduced to

FIG. 96. Branching thorn of the Honey-Locust (Gleditschia), an indurated branch
developed from an accessory bud produced above the axil. a. Three buds under the
base of the leaf-stalk, brought to view in a section of the stem and leaf-stalk below.

FIG. 97. Thorn of the Cockspur Thorn, developed from the central of three axillary
buds; one of the lateral buds is seen at its base.

SPECIAL FORMS.

ay
t

7

seales , and the advancing apex rises at length into an ordinary
stem, while the opposite and older end gradually dies away. A
bud forms in the axil of each seale-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 hoe or plough merely hastens the
establishment of as many new plants, each with
roots, bud, and a small store of nourishment
ready provided. It is this which makes Couch-
Grass or Quick-Grass (Triticum repens) very

a

ees

98

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, ¢. 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, being 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 from the
under side or the whole surface. Thus established, the older

FIG. 98. Slender rhizoma of Carex arenaria, of Europe, which binds shifting sands
of the sea-shore.

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

‘FIG. 99. A piece of the rootstock of the Peppermint, enlarged, with its node or joint,
and two axillary buds ready to grow.

58 MORPHOLOGY OF STEMS.

portions die and decay as corresponding additions are made to
the opposite growing extremity. -Each year’s growth is often

marked conspicuously, sometimes by a strong contraction where
the interruption took place, as in certain species of Iris (Fig. 216);

or by the circular im-

pressed scar (likened to

the impression ofa seal)
* jin Solomon’s Seal; this
ye, being the place where
J the annual aerial stem,
bearing the vegetation,
separated in autumn
from the perennial rhi-
zoma. The numerous
slender lines encircling the rootstock are the scars left after the
decay of the scale-like leaves or bud-seales, such as are seen at
4 the young and growing end of the rootstock.
The rootstock of Diphylleia, of the Alleghany
Mountains (Fig. 101), is similar; but the
yearly growths are so exceedingly short that
they become vertical, the bud of each year,
is close to the stalk of the year preceding,
and the scarsanarking preyious growths are
in contact.? villi makes a short and
mostly vertical rootstock, which, when it
remains simple and dies away promptly
below (as in Fig. 102), comes nearly within
the definition of a corm. But in several

1 The rootstock in Polygonatum and Diphylleia is a sympodium (110, note),
the terminal bud developing yearly the growth above ground and perishing

FIG, 100. Rootstock of Polygonafum or Solomon’s Seal, with the terminal bud, the
base of the stalk of the season, and three scars from which the latter has separated in
as many former years.

FIG. 101. Rhizoma of Diphylleia eymosa, showing six years’ growth, and a bud for
the seventh: a, the bud: 6, base of the stalk of the current year: c, scar left by the ©
decay of the annual stalk of the year before; and beyond are the scars of previous years. —

FIG. 102. Shoot and young rootstock of Trillium erectum, with only terminal bud.

3

9

ube

SPECIAL FORMS.

species, and in older individuals, it is longer, often oblique, and
branching, and bears the scars from which the annual aerial
growths have separated.*/ Nymphzea odorata,
the sweet-scented white Water Lily, grows by
very long, stout, and simple rootaiagl es: ) in
NW; tuberosa the sides of the. rootstag produce
short lateral branches or tubers.

117. A Tuber may be morphologically char-
‘acterized as a short thickened rhizoma on a
slender base, or a rootstock some portion of
which —mostly a terminal portion and involy-
ing several nodes —is thickened by the depo-
sition of nourishing matter. A potato and a
Jerusalem artichoke are typical examples
(Fig. 104-107) ; and the difference between
these subterranean branches and the roots which they may bear
is very obvious. Their eyes are axillary buds; the leaves which
subtend them are plainly dis-
cernible, in the form of short and
closely appressed scales. In the
attempt, occasionally seen, to
form axillary tubers above-
ground by the Potato-plant, the
leafy nature of the scales is
evidenced. (Fig.
105.) By heaping
‘the soil around
the stems, the
number of tuber- B
iferous branches
may be __ in-
creased. The number of nodes and internodes involvea in a
tuber may be many or few. There is one instance of what may

in autumn, to be renewed by an axillary bud, which makes its subterranean
growth and the rudiments of the aerial in early summer.
1 This rhizoma is a monopodium, being continued year by year by the
terminal bud, and the aerial stem or stems sent up in spring, bearing the
’ whorl of leaves and blossom, are axillary branches.

FIG. 103. An older and longer one of the same spec ies, showing branches. scars left
by former leaf- and flower-bearing stems: also at tip (stripped of the covering scales), the
bases of two such stems of the season, and the terminal bud between them, for the con-
tinuation of the growth of the rootstock, &c., the next season.

FIG. 104. Base of stem of Helianthus tuberosus, or Jerusalem Artichoke, developed
from a tuber, and producing a second generation of tubers.

FIG. 105. Monstrosity of a Potato-plant, with an axillary bud developing into some-
thing between a bulblet and a tuber; the scales represented by obvious leaves. (From
the Gardeners’ Chronicle. ) ~

a
.

GU MORPHOLOGY OF STEMS.

be called a Monomerous tuber, namely in Nelumbium luteum
(Fig. 1087), where it consists of a single thickened internode of

107 106 108

an aquatic runner, which is accordingly quite destitute of scales
or buds. ‘The growth proceeding from this simple tuber is
necessarily from a bud of
the node at its apex, whence
also a. cluster of roots is
produced. Of a somewhat
similar nature are the con-
catenate tubers of Apios
tuberosa (several of which
1037 are strung as it were upon’
a long filiform axis), the tubers not unfrequently being mo-
nomerous, although the larger ones are not so.
117°. Tubereles, as they may be termed, are of a a or
ambiguous character between tubers and tuberous roots. A good
example of the latter is afforded by Dahlia-roots. (Fig. 68.)
They yield their nourishing substance to growing buds on the
stem above, but do not themselves normally produce even

FIG. 106. Forming potatoes in various stages. 107. One of the younger ones en-
larged. 108. Section of a small portion passing throngh an eye, or bud, more enlarged.

FIG. 108¢, A monomerous (i. e. one-membered) tuber of Nelumbium luteum, formed
of a single internode.

SPECIAL FORMS. 61

adventitious buds. Sweet potatoes (55), although equally
roots, do produce adventitious buds, especially from near the
upper end. The somewhat similar tubercles or tumefied roots
of certain Orchises and other plants of the same tribe,! definite
in number and shape, and sometimes imitating a corm, are
charged 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 tumefied short root from its
very base.”

118. A Corm (Cormus) is to be compared on the one hand
with a short rootstock or tuber, on the other with a bulb. It
is a subterranean fleshy stem, of rounded or depressed figure
and solid texture. Some of its buds grow into new corms, and
these, upon the death of or separation from the parent, become
new individuals: some develop above ground alla. I
the vegetation and the blossoms of the season. if KS i z
A good type of corm is that of Cyclamen LAN
(Fig. 109), in which the very base of the
seedling stem grows fleshy, and widens from
year to year, but hardly at all lengthens, and ;
so becomes far broader than high, or de-
pressed. As the main bulk belongs to the 109
first internode, or caulicle, the buds from which the yearly
growths of leaves and flower-stalks spring are at the centre of
the summit or upper surface, the roots
from the lower, and the sides seldom pro-
duce any buds. The corm of Indian
Turnip (Ariszema triphyllum, Fig. 110)
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 corms.
¢ 119. But in Crocus (Fig. 111, 112), Colchicum (Fig. 117),
Gladiolus, and the like, the sheathing bases of one or two leaves
enclose the corm with a membranous-scaly coat, giving it exactly
the appearance externally of a coated bulb. Such have been not
inappropriately named solid bu/bs. Inecommon parlance, they
will doubtless continue to be called bulbs, and even in popular

1 Not, however, such as those of Aplectrum, Tipularia, etc., which are
genuine corms or tubers. =

* Irmisch, Beitr. Biol. & Morphol. Orchid. 1853, fide Duchartre, Elém.
Bot. 278.

FIG. 109. Depressed corm of Cyclamen.
FIG. 110. Corm of Indian Turnip, Arisema triphyllum.

62 MORPHOLOGY OF STEMS.

botanical descriptions. In fact, while they differ from naked
corms in having some investment, they differ from true bulbs
only in the greater size of the solid axis and the fewness of
the investing scales;

the stem or solid body
making the greater part
of the corm, but a very
small part of a proper
bulb. There are, more-
Ny, over, all gradations be-
11 112 tween the two.

120. A Bulb, as compared with a corm, may be said to be 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
and thickened leaves or the thickened bases of ordinary leayes.
Compared with buds (73), it is a very fleshy bud, usually large
and subterranean, the axis of which never elongates. It is a

provision for future growth, the stored nourishment of which is
deposited in the leaves, or the homologues of leaves, instead of
in the stem.

FIG. 111. Corm of Crocus, théfew thin enveloping scales removed, showing their
scars, which mark the nodes, the shrivelled vestige of the last year’s corm at the base,
and buds developing into new ones on various parts of its surface. 112. Vertical section
of a similar corm, with a terminal and one lateral bud.

FIG. 113. Section of a tunicated bulb of the Onion.

FIG. 114. Vertical section of the bulb of the Tulip, showing its stem or terminal
bud (c) and two axillary buds (8, 5).

FIG. 115. Bulb of a Garli:, with a crop of young bulbs.

FIG. 116. Vertical section of the corm of a Crocus: a. new buds.

FIG. 117. Vertical section of the corm of Colchicum (6), with the withered corm of
the preceding (a), and the forming one (c) for the ensuing year.

SPECIAL FORMS. 63

121. A Tunicated or Coated Bulb (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 Sealy Bulb
has the bulb-scales
comparatively —nar-
row, thick, and small,
imbricated, but not
severally enwrapping
each other. That of
the Lily is the mest |
familiar and char- Wd)
acteristic example.
Mie) 118; 119:)

123. Bulblets are small aerial bulbs, or buds with fleshy scales,
which arise in the axils of the leaves of several plants, such as
the common Lilium bulbiferum and
L. tigrinum, the Tiger Lilies of the
gardens (Fig. 120). Here they 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 bulblets
take the place of flower-buds in the
umbel. Bulblets plainly show the identity of bulbs with buds.

124. All these extraordinary, no less than the ordinary, forms
of the stem, grow and branch, or multiply, by the development
of terminal and axillary buds. This is perfectly evident in the
rhizoma and tuber, and is equally the case in the corm and bulb.
The stem of the bulb is usually reduced to a mere plate (Fig.
114 4a), which produces roots from its lower surface, and leaves
or scales from the upper. Besides the terminal bud (c), which
usually forms the flower-stem, lateral buds (4, 6) 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 become bulbs, feeding on the parent bulb,
which in this way is often consumed by its own offspring, as in

=)

FIG. 118. Sealy bulb of Canada Lily, Lilium Canadense, after flowering. 119. Ver-
tical section of same, showing two new young bulbs within.
FIG. 120. Bulblets in the axil of the cauline leaves of Tiger Lily.

64+ MORPHOLOGY OF STEMS.

the Garlic (Fig. 115); or, finally separating from the living
parent, just as the bulblets of the Tiger Lily fall from the stem,
they may form so many independent individuals. So the corm
of the Crocus (Fig. 111, 112) produces one or more new ones,
which feed upon and exhaust it, and take its place; and the
next season the shrivelled remains of the old corm may be found
underneath the new. The corm of Colchicum (Fig. 117) pro-
duces a new bud on one side at the base, and is consumed by it
in the course of the season; the new one, after flowering by its
terminal bud, is in turn consumed by its own offspring; and
so on. The figure represents at one view, a, the dead and
shrivelled corm of the year preceding; 6, that of the present
season (in a vertical section); and, ¢, the nascent bud for the
growth of the ensuing year.

125. Condensed Stems, homologous with corms, tubers, &e.,
and similar in mode of growth, but above ground, and multiply-
ing in the same ways, are not uncommon. The Cactus family is
mainly composed of such forms, of flat- or round-jointed Prickly
Pears (Opuntia), fluted or angled columns (Cereus), and glob-
ular Melon-Cactus, Mamillaria, and Echinocactus. The latter
types, which completely imitate corms, are the most consolidated
forms of vegetation. While ordinary plants are constructed on
the plan of great expansion of surface, these present the least
possible amount of surface in proportion to their bulk, their
permanent spherical figure being that which exposes the smallest
portion of their substance to the air. Such plants are evidently
adapted to very dry regions ; and in such only are they naturally
found. Similarly, bulbous and corm-bearing plants, and the
like, are a form of vegetation which in the growing season may
in the foliage expand a large surface to the air and light, while
during the period of rest the living vegetable is reduced to a
globular or other form of the least surface ; and this is protected
by its outer coats of dead and dry scales, as well as by its subter-
ranean situation ;— thus exhibiting another and very similar
adaptation to a season of drought. And such plants mainly
belong to countries (such as Southern 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 beneath being early cut off by drought, the plants
rest securely in the corm-like forms to which they are reduced,
and retain their moisture with great tenacity until the rainy season
returns. Then they shoot forth leaves and flowers with wonderful
rapidity, and what was perhaps a desert of arid sand becomes
green with foiiage and gay with blossoms, almost in a day.

SPECLAL FORMS. 65

126. Stems serving 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 fleshy 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 Bossiza and Carmichelia among Legu-
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 (7. e. wooden-
leaved) section of Phyllanthus, and in Phyllo-
cladus (New Zealand and Tasmanian trees of
the Yew family). In all these, the cauline nature
is manifest by the continuous or proliferous
growth, by the marked nodes and internodes,
and often by the bearing of flowers.

127. Cladophylla (literally, branch-leaves) are more ambigu-
ous in character. The most familiar examples are found in the
peculiar foliage of Ruscus, Myrsiphyllum, Asparagus, and in
some other genera of the same family. In these the primary or
proper leaves of the shoots are little scales, one to each node, and
quite functionless. From the axil of each is immediately pro-
duced a body answering in all respects to the blade of a leaf,
both in appearance and in office. They also accord with leaves
in being expanded horizontally, although they take a twist which
brings them more or less into a vertical position, in the manner
of phyllodia (that is, of leaf*stalks assuining the form and office
of leaf-blades, 217); wherefore they may be regarded as the
first and only leaf of an axillary branch with the internode
under the leaf wholly undeveloped and no further growth ever
taking place. But, on the other hand, their anatomical structure
is said to be that of stems rather than of leaves. Moreover,

FIG. 121. Foliiform branch of Mublenbeckia platyclada. growing from the apex,
bearing a smal! and transient leaf at some nodes, also a fluwer or two.

66 MORPHOLOGY OF STEMS.

the cladophyll of Ruscus (called Butcher’s Broom in England,
Fig. 123) not only becomes firm, hard, and spiny-tipped, but
it exhibits the character of a
branch by bearing flowers on the
middle of one face, in the axil of
a little bract. Under this view
such a cladophyll would seem to
be a flattened branch of two in-
ternodes, or else of one internode
with a flower-stalk adnate to it.
In Myrsiphyllum (a South A fri-
can climber, commonly cultivated
under the erroneous name of
Smilax, Fig. 122), the cladophyll
is wholly leaf-like in appearance
as well as in function, and it never bears either scale-leaf
or blossom; but the flowers are on slender stalks from buds out
of the same axil. (See Dickson in Trans. Bot. Soc. Edinb.
xvi., and Van Tieghem, Bull. Bot. Soc. France, xxxi., for a
discussion of the nature of cladophylla.)

128. To all such leaves or imitations of leaves, Bischoff has
given the name PHYLLocLapiA, sing. PHytLoctapium. To those
definitely restricted to one internode, and which so closely
counterfeit leaves, Kunth gave the name of CLapopra, sing.
Ciapopium. 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 imitating the blade of a leaf. ‘The
name Phyllocladium (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, Kunth’s name of Cladodium (t. e. a
branch-like body) is false in meaning, and may
be replaced by that of CLApopiyLLuM (7. e. leaf-
branch), or in shorter English CLapopnyLt.

129. Frondose Stems. Finally, in some few
124 pheenogamous plants, the whole vegetation is re-
duced toa simple leaf-like expansion, as in Duckweed (Lemna),

122 123

FIG. 122. Myrsiphyllum, with cladophylls serving for foliage; the true leaves eon-
sisting of minute and very inconspicuous scales subtending the former.

FIG, 123. Asingle cladophyll of Ruscus aculeatus in the axil of a scale-leaf, bearing
another scale-leaf on the middle of its face, and flowers in the axil of this.

FIG. 124. Lemna minor, a common Duckweed, whole plant in flower, magnified.

INTERNAL STRUCTURE. 67

Fig. 124. Here is no differentiation whatever into stem and
foliage ; but the expanded floating body which serves for both
must be counted as stem developed horizontally into a flat plate,
for it produces a root from the under surface and a flower from
the edge. This simplification is common in some orders of
Cryptogamous plants ; and such a body, which answers both for
stem and foliage, is termed a Fronp, from the Latin frons, which
means either leaf or leafy bough. In some species of Lemna
the frond is thickened or plano-convex : in Wolffia, the simplest
and smallest of phanogamous plants, it is a globular green mass,
seldom much larger than the head of a pin, wholly destitute of
root, propagated by proliferous budding from one side, and from
within the top producing a flower or pair of flowers.

§ 3. INTERNAL STRUCTURE.

130. The investigation of the intimate structure of the stem,
as of the other organs, belongs to vegetable anatomy or histology
(treated in Part II.) ; but the general outlines of structure, so
far as is requisite to the explanation of what is visible to the
naked eye, should be here explained.

13i. The stems of phzenogamous plants anatomically consist
of two general elements, the cellular and the woody ; the former
exemplified in the commoner stems by the pith and outer bark, the
latter by the wood. Bothare equally composed of cells, or origi-

cs

a A
Hat oa ae

124%
nate as such ; but those which form the woody system of the stem
mainly undergo, 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 vessels. ‘The latter are almost

Jee Se EE EE ee eee eee ee eee eee

PIG. 124%. A magnified slice of a portion of the flower-stalk of Richardia ZEthiopica
(the so-called Calla Lily), transverse with some longitudinal view: mainly parenchyma,
the vells built up so as to leave comparatively large vacancies (intercellular spaces or
alr-passages); near the centre a cross-section of a fibro-vascular bundle, and next the
margin or rind some finer ones.

638 MORPHOLOGY OF STEMS.

always associated with 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-vascular bundles. These
run lengthwise through the stem, sometimes as such separate
threads, sometimes confluent into a compact structure. The
softer or at least the non-fibrous portions, formed of comparatively

a b d

000 uo9

0) O00
0

a>

092 920 QO

VON 0

ung

WOOUOCD.O0 0 6

hw)

Uo

CCK

=
EC
Zg

(Cauca
((

Cac

LIC

c
125

short and commonly thin-walled cells, form cellular tissue. Its
ordinary form (of roundish, cubical, or polyhedral and thin-walled
cells) is called parenchyma. This abounds in herbaceous stems or
herbaceous parts: in trees and shrubs, woody tissue largely pre-
vails; in most herbs, it forms a notable portion; in some (especially

FIG. 125. Fibro-vyasecular elements. a. Bast-cells (long wood-cells) of fibrous bark
of Linden or Bass-wood. b. Some wood-cells and (below) a duct, and ¢ a detached
wood-cell of the wood of same tree, equally magnified with a. d. A detached wood-
cell from a shaving of White Pine, showing the peculiar disk-like markings. e. Portion

of same shaving. jf. Portion of a dotted duct from the Vine, evidently made up of a
series of short cells. g. Part of a smaller dotted duct, showing no appearance of such
eomposition. hk, i. Spiral ducts or vessels, of the ordinary kind. j. Spiral duct of
Banana. k. Duct from Celery, the thread within spiral or annular below, reticulated
above, and higher passing into the state of dotted duct. /. Duct from Impatiens, with
the open spiral passing into rings at the middle. All magnified somewhat equally.

INTERNAL STRUCTURE. 69

in certain aquatic herbs), it is reduced to a few threads or
vessels, generally delicate, and sometimes obscure. The ac-
companying 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 parenchyma ; but
an early differentiation takes place, converting certain portions
into woody or fibro-vascular tissue. This is arranged in two
ways, giving rise to two kinds of stem in phenogamous plants,
which haye been termed the Endogenous and the Exogenous,’
meaning inside and outside growers.

133. The two plans of stem are usually manifested in external
conformation as well as internal structure, and are correlated
with important differences in embryo, foliage, and flower.” Palms,
Lilies, 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, are of the exogenous class.
In an exogenous stem, the wood occupies annual concentric layers,
one of each year’s growth; the centre is occupied by a pith,
composed of parenchyma only, the circumference by a separable
bark; so that a cross-section presents a
series of rings or circles of wood, or in
the first year one ring, surrounding the §
pith and surrounded by the bark. An jj)
endogenous stem has the wood in distinct : i i
threads or fibro-yascular bundles, travers- § | | |
ing the cellular system or parenchyma with 9) | |
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 Terms introduced by DeCandolle, following the ideas of Desfontaines,
and which have played an important part in structural and systematic
botany ever since DeCandolle adopted these names as those of the two
primary divisions of phenogamous plants, Lxogene and Endogene. But it
has long been seen that the name of the second kind is not appropriate; and
the older and better (though longer) names of Jussieu, Monocoiyledones and
Dicotyledones, are reverted to. Yet the Candollean names are still much
employed, with due explanation, to designate the two kinds of structure of
the stem.

2 Yet with some more or less valid exceptions, as when the annual stem
of Podophyllum and the rhizoma of Nymphza, among dicotyledonous plants,
imitate the endogenous structure; or where the pith of an evidently exogenous
stem, as in the Piperacew, has scattered woody bundles in an endogenous
fashion ; or where monocotyledonous plants have all their woody bundles in
a definite circle, as in Luzula, Croomia, &c.

FIG. 126. Se tion of a small Palm-stem, in two directions.

70 MORPHOLOGY OF STEMS.

the whole section, or when few in number of somewhat definite
position or arrangement. The ordinary appearance of such a
stem, both on the longitudinal and the cross-section, is shown
in Fig. 126; it may also be examined in the Cane or Rattan,
the Bamboo, and in the annual stalk of Indian Corn or of
Asparagus. The appearance of ordinary wood is very familiar.

135. The newer woody bundles of an endogenous stem are yari-
ously intermingled with the old. When DeCandolle gave the name,
it was supposed, from Desfontaines’s researches, that the older
bundles occupied, or came at length to occupy, the circumference
of the trunk, while only new ones were formed in the centre ;
and that increase in diameter, when it took place at all, resulted
from the gradual growth and distention of the whole. Hence
the contrasting name of endogenous, or inside growing, and for
such plants the name of ENpoGENous PLAnts, or ENDOGENS.
Our actual knowledge of the structure and growth of these stems,
as will be seen, cannot be harmonized with this view in any
way which gives to the name endogenous an appropriate signifi-
cation. The name continues as a counterpart to the more correct
one of exogenous, and as a survival of former ideas.

136. The Endogenous Structure (so called) of the stem is cor-
related with a monocotyledonous embryo (39), usually with a
ternary arrangement in the flower (322), and commonly with
parallel-veined leaves. (173.) Endogens, although they have
many herbaceous and a few somewhat woody representatives
in cool temperate climes, mostly attain their full variety of fea-
tures and rise to noble arborescent forms under a_ tropical
sun. Yet Palms —the arboreous type of the class — do extend
as far north in this country as the coast of North Carolina (the
natural limit of the Palmetto, Fig. 126") ; while in Europe the
Date and the Chamerops thrive in the warmest parts of the Euro-
pean shore of the Mediterranean. The manner of their growth
gives them a striking appearance ; their trunks being unbranched
cylindrical columns, rising to the height of from thirty to one
hundred and fifty feet, and crowned at the summit with a simple
cluster of peculiar foliage. Palms generally grow from the
terminal bud alone, and perish if this bud be destroyed; they
erow 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 ordinary species, regularly
in the Doum Palm of Upper Egypt, and in the Pandanus, or
Screw-Pine (Fig. 69), which belongs toa family allied to Palms :
in such cases the branches are cylindrical. But when lateral

INTERNAL STRUCTURE. 71

buds are freely developed (as in the Asparagus), or the leaves
are scattered along the stem or branches by the full development
of internodes (as in
the Bamboo, Maize,
&ce.), they gradually
taper upward in the
manner of most ex-
ogenous stems.

for. “This~ kind
of stem comprises
several subordinate
types as to internal

('X

ry aN SP =

: a
structure, which to a
be well understood ba
must be studied his- a
tologically, under the Ea

ae
Hf

pase

microscope.’ Toone
of these, by no means
the simplest, belongs
the ordinary palm-
stem, the anatomy
of which was made

classical by Mobhl, 3
and has been 5 3

wi ¢
supplemented \ N\ Ay , :
by Negeli. 9 ey WZ i
In this a large = \lf \ Ki

part of the =Zsthy
bundles, or all 7).

of the more
conspicuous kind, starting from the base of the leaf to which
they respectively belong, curve 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 accessible memoir on the subject, of recent date,
see Guillaud, Recherches sur l’Anatomie comparée et le Développement des
Tissus de la Tige dans les Monocotylédones, published in Ann. Sci. Nat.
ser. 6, v. 1-176, 1877. Six types of the stem of Monocotyledons are here
recognized by anatomical characters and modes of growth, one of them
having four modifications.

FIG. 126+ Sabal Palmetto in various stages; also the Yucca aloifolia or Spanish
Bayonet.

72 MORPHOLOGY OF STEMS.

as shown in Fig. 127. It is partly owing to this connection of
these fibres with the rind that the latter is not separable from
the stem. In some Palms, and in Grasses,
there is no marked distinction between
4 the wood and rind, or no proper rind at
Hall. In others, such as the Palmetto
4 (Fig. 126), there is a marked rind
or false bark, which receives independent
fibro-vascular bundles from the leaf-stalks,
and is traversed by them in parallel lines.
oq In Grass-stems, and others with long inter-
ib i nodes and closed nodes, the fibro-vascular
C/iis bundles all run approximately straight
: and parallel through the internodes, but are
jij intricate and anastomosed in the nodes.
5d The whole centre of the internodes, when
not hollow or before it becomes so, is occupied by a true pith,
like that of an Exogen, and in some cases equally destitute of
fibro-vascular bundles, but often with scattered ones, after the
manner of certain Exogens anomalous in this respect, such as
Nyctaginaceze and some Araliacee. Endogenous stems of
simpler structure, as in herbaceous Liliaceze, Commelynacez,
&c., have a distinct cortical portion (at least in the root-stock
or-portion of stem properly comparable with palm-trunks and
the like) ; but this is mostly destitute of fibro-vascular bundles.
Most of them have two kinds of vascular bundles, one
of which 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 within this circle, either of
very few and scattered bundles, as in Convallaria majalis, or
numerous and scattered, as in Uvularia and the leafy stems of
Tradescantia Virginica ; or these bundles are few and arranged
nearly in an inner circle close around 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, dividing a purely cellular medullary 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 until it attains its limit. Ligneous and enduring
stems increase similarly up to a certain period. Then the rind

WN ll

{ )

FIG. 127. Diagrammatic view of the curved course of the fibro-vascular bundles

in a palm-trunk.

INTERNAL STRUCTURE. 73

sooner or later ceases to distend or adapt itself to further in-
crease in diameter, and there is no interior provision for indefinite
increase in the greater number of woody endogenous trunks. But
in Draczena (Dragon-trees), in the arborescent Yuccas, and the
like, the zone intermediate 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 haye done) ; and they imitate exogenous trunks
to a considerable extent in mode of growth.

139. The wood of an endogenous woody stem is hardest and
most compact at the circumference; in palm-stems commonly
it is largely mixed with parenchyma or pith at the centre, even
in old trunks.

140. The Exogenous Structure, that of ordinary wood, is char-
acterized by the formation of a distinct zone of wood between a
central cellular medullary portion (pith) and an outer chiefly
cellular portion (bark), traversed by plates from the pith (medul-
lary 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, f-/), at definite points
in the common parenchyma of the initial stem (four equidistant.
ones in the Sugar Maple) ; each is soon surrounded by incipient

proper wood-cells (Fig. 125, b, c), together forming a fibro-vascular
bundle or thread. Additional ones are intercalated as the second
and third internodes develop, and so a column (in cross-
section a ring) of wood is produced, always so arranged as to

FIG. 128. Diagram of a cross-section of a forming seedling stem, showing the
manner in which the young wood is arranged in the cellular system.

FIG. 129. The same at a later period, the woody bundles increased so as nearly to
fill the circle.

FIG. 130. The same at the close of the season, where the wood has formed a com-
plete circle, interrupted only by the medullary rays, which radiate from the pith to
the bark.

74 MORPHOLOGY OF STEMS.

surround a purely cellular central part (the pith), while sur-
rounded by a cellular external rind, the bark, or outer bark.
The diagrams (Fig. 128-150) rudely show some stages in the
formation of the zone of wood. The fibro-vascular bundles
originate in the bases of the leaves, and develop outward into the
forming leaves as well as downward ifto the forming stem.

142. First Year’s Growth. The wood, even in a herbaceous
or annual stem, at the completion of the first year’s growth,
forms a zone or tube,
enclosing the pith. But
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 divide the wood
into wedges. These
lines, forming what is
called the silver-grain
in wood, are the Mrp-
ULLARY Rays. They
represent the cellular
system of the wood it-
self, or untransformed
parenchyma. Being
pressed by the woody

of

PUN CCU CHP CHECK Cer eLE

tt

EF,
ssa

RECRETTC TCC

LK
Pe wedges, their cells are
Rene cit va e «¢ aif @ pe laterally fattemedsssam
38 some stems, the med-

ullary rays, or many
of them, are comparatively broad and conspicuous; 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 they usually continue to grow through the
season ; that, is the wood grows externally. The general ana-

FIG. 131. Longitudinal and transverse section of a stem of the Soft Maple (Acer
dasycarpum), at the close of the first year’s growth; of the natural size.

FIG. 132. Portion of the same, magnified, showing the cellular pith, surrounded by
the wood, and that by the bark.

FIG. 133. More magnified slice of the same, reaching from the bark to the pith:
a. part of the pith; 0. vessels of the medullary sheath; ec. the wood; d, d. dotted ducts in
the wood; ¢, e. annular ducts; 7. the liber, or inner fibrous bark; g. the cellular envelope,
or green bark; h. the corky envelope; 7. the skin or epidermis; &. one of the medullary
rays, seen on the transverse section.

INTERNAL STRUCTURE. i)
tomical structure of a woody exogenous stem of a year old is
displayed in the Fig. 131-135. Viewing the parts particularly,
and in order from centre to circumference, there is, —

Ist. 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 Conifers, the wood is of uniform structure, being
wholly composed of a woody tissue with peculiar markings (Fig.
125, d,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, 7. 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 immediately surrounding the pith, they have
collectively been termed the medullary sheath, but they hardly
deserve a special name. ‘The vertical section in Fig. 153 divides
one of the woody wedges, and shows no medullary ray ; but there
is one at the posterior edge of the transverse section. But, in the
much more diagramatic Fig. 154, 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 (4).
These medullary rays form the silver-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. 155, 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-vascular threads are found dispersed
through the pith, presenting a somewhat remarkable anomaly. This
occurs in Aralia racemosa, and more strikingly in Mirabilis and other
Nyctaginacee, and in Piperacexw. (133, foot-note.)

76 MORPHOLOGY OF STEMS.

tal system of the wood, or the woof, into which the vertical woody
fibre, &c., or warp, isinterwoyen. The inspection of a piece of oak
or maple wood at once shows the pertinency of this illustration.

i

hel

ti
i

135

3rd. The Bark or rind. This at first consisted of simple
parenchyma, like that of the pith, except for the green color
developed in it, the same as that which gives verdancy to foliage.
This green matter is formed in the cells of all such parts when
exposed to light, consists of green grains of somewhat complex
chemical composition, has important functions to perform in
assimilation (¢.e. in the conversion of the plant’s crude food into
vegetable matter), and is named CHLOROPHYLL, 7. e. leaf-green.
The completed bark, when all its parts are apparent, as espe-
cially in-most trees and shrubs, is composed of three strata, of
which the green bark, the most conspicuous in the young shoot,
is the middle layer, therefore named the MesopnHtaum. This is
soon covered, and the green color obscured, by a 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 CorKxy
EnvVeELore or layer, or EpipHta@um. The latter name denotes its
external position; the former, that it is the layer which, when
much developed, forms the cork of Cork-Oak and those corky
expansions which are so conspicuous on the twigs of the Sweet
Gum (Liquidambar), and on some of our Elms (Ulmus alata
and racemosa). It also forms the paper-like exfoliating layers
of Birch-bark. It is composed of laterally flattened parenchy-
matous cells, much like those of the Eprpermuis (Fig. 133, 2),
which directly overlies it, and forms the skin or surface of the

FIG. 134. Vertical section through the wood of a branch of the Maple, a year old,
so as to show one of the medullary rays, passing transversely from the pith (p) to the
bark (6): magnified. But a section can seldom be made so as to show one unbroken
plate stretching across the wood, as in this instance.

FIG. 135. A vertical section across the eucs of the medullary rays: magnified.

T7

INTERNAL STRUCTURE.

stem, and of the whole plant. Lastly, the inner bark, accord-
ingly named EnporHievm, takes the special name of Liser, and
is the most important portion of the bark in the stems of trees
and shrubs. Complete and well-developed liber, like that of

Linden or Basswood, contains two peculiar kinds of cells in
BS C Ww

cea)

QLO<e
tienes

aan
oa
50
vs)

1354 mr
addition to common parenchyma, both of the fibrous or vascular
class: viz., 1. Criprirorm or StevE-cELLs, a sort of ducts the
walls of which have open slits, through which they communicate
with each other; 2. Bast or Bast-cexis, the fibre-like cells
which give to the kinds of inner bark that largely contain them

FIG. 135¢ Portion of a transverse section (above), and a corresponding vertical see-
tion (below), magnified. reaching from the pith (p) to the epidermis (e) of a stem of
Negundo, a year old: B. the bark; W. the wood; and C. the cambium-layer, as found
in February. The parts referred to by small letters are: p.a portion of the pith:
mr.small portion of a medullary ray where it runs into the pith; four complete med-
ullary rays as seen on a transverse section, appear in the upper figure, running from
pith to bark: ms. medullary sheath, a circle of spiral unrollable ducts, one seen length-
wise with uncoiling extremity in the lower figure: w, w. woody tissue: dd. one of the
dotted ducts interspersed in the wood: c/. carnbium-layer or zone of new growth of
wood and inner bark: /-b. liber or inner bark, the inner portion of which is here cellular,
the outer (6) composed of slender and thick-walled bast-cells or true liber-cells:
ge. green envelope or inner cellular bark: ce. corky envelope or outer cellular bark:
e. epidermis.

(9 6)

7 MORPHOLOGY OF STEMS.
their strength and toughness. They are like wood-cells except
in their greater length and flexibility, and in the thickness of
their walls, which greatly exceeds the calibre. This is the
material which gives to the bast or inner bark of Basswood, &c.,
the strength and pliability that adapts it for cordage and for
making mats: it is the material of linen, and the like textile
fibres. (For a view of the whole composition and structure of a
woody stem at the close of the first year’s growth, and immedi-
ately before that of the second year begins, see Fig. 135%.)

143. Annual Increase in Diameter. An herbaceous stem does
not essentially differ from a woody one of the same age, except
that the wood forms a less compact or thinner zone; ‘and the
whole perishes, at least down to the ground, at the close of the
season. Buta woody stem makes provision for continuing its
growth from year to year. As the layer of wood continues to
increase in thickness throughout the season, by the multiplication
of cells on its outer surface, between it and the bark, and when
growth ceases this process of cell-multiplication is merely sus-
pended, so there is always a zone of delicate young cells in-
terposed between the wood and the bark. This is called the
Cambium, or, better, the Camprum-Layer. It is charged with
organizable matter, which is particularly abundant and mucila-
ginous in spring when growth recommences. This mucilaginous
matter was named Cambium by the older botanists: they sup-
posed — as is still popularly thought — that the bark, then so
readily separable, really separated from the wood in spring, that a
quantity of rich mucilaginous sap was poured out heowern them
and became organized into a tissue, the inner part becoming new
wood, the outer, new bark. But delicate slices show that there ~
is then no more interruption of the wood and inner bark than
at any other season. The bark, indeed, is then very readily
detached from the wood, because the cambium-layer is gorged
with sap; but such separation is effected by the rending of a
delicate forming tissue. And if some of this apparent mucilage
be scraped off from the surface of the wood, and examined under
a good microscope, it will be seen to be a thin stratum of young
wood-cells, with the ends of medullary rays here and there in-
terspersed. The inner portion of the cambium-layer is therefore
nascent wood, and the outer is nascent bark. As the cells of
this layer multiply, the greater number lengthen vertically into
woody tissue: some are transformed into ducts; and others,
remaining as parenchyma, continue the medullary rays or com-
mence new ones. In this way, a second layer of w ood is formed
the second season over the whole surface of the former layer

INTERNAL STRUCTURE. 79

between it and the bark; and this is continuous with the woody
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 occupying 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. Demareation of the Annual Layers results from two or more
causes, separate or combined. In oak and chestnut wood, and
the like, the layers are strongly defined by reason of the accumu-
lation of the large dotted ducts (here of extreme size and in
great abundance) in the inner portion of each layer, where their
open mouths on the cross-section are conspicuous to the naked
eye, 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 are most distinct in trees of temperate climates like
ours, where there is a prolonged period of total repose, from the winter’s
cold, followed by a vigorous resumption of vegetation in spring. In tropical
trees, they are rarely so well defined; but even in these there is generally a
more or less marked annual suspension of vegetation, occurring, however,
in the dry and hotter, rather than in the cooler season. There are numerous
cases, moreover, in which the wood forms a uniform stratum, whatever be
the age of the trunk, as in the arborescent species of Cactus; or where the
layers are few and by no means corresponding with the age of the trunk, as
in the Cycas.

In many woody climbing or twining stems, such as those of Clematis,
Aristolochia 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-
posed, passing at right angles to each other from the circumference towards
the centre, so that the transverse section of the wood nearly resembles a
Maltese cross. But these are exceptional cases, which scarcely require
notice in a general view.

80 MORPHOLOGY OF STEMS.

vigorous vernal growth, with much larger cells, thus marking
an abrupt transition from one layer to the next. Besides being
finer, the later wood-cells of the season are commonly flattened
antero-posteriorly, probably by growing under greater pressure.

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

146. Sap-wood (Atsurnum). In the plantlet and in the
developing bud, the sap ascends through the whole tissue, of
whatever sort: at first through the parenchyma, for there is then
no other tissue; and the transmission is continued through it,
especially through its central portion, or the pith, in the growing
apex of the stem throughout. But, in the older parts below, the
pith, soon drained of sap, becomes filled with air in its place,
and thenceforth it bears no part in the plant’s nourishment. As
soon as wood-cells and ducts are formed, they take an active
part in the conveyance of sap, for which their tubular and ca-
pillary character is especially adapted. But, the ducts in older
parts, except when gorged with sap, contain air alone; and in
woody trunks the sap continues to rise year after year to the
places where growth is going on, mainly through the proper
woody tissue of the wood. In this transmission, the new layers
are most active ; and these are in direct communication with the
new roots on the one hand and with the buds or shoots and leaves
of the season on the other. So, by the formation of new annual
layers outside of them, the older ones are each year removed a
step farther from the region of growth; or rather the growing
stratum, which connects the fresh rootlets that imbibe with the
foliage that elaborates the sap, is each year removed farther from
them. The latter, therefore, after a few years, cease to convey
sap, as they have long before ceased to take part in any vital
operations. The cells of the older layers, also, usually come to
have thicker walls and smaller calibre than those of the newer.
Thus arises a distinction — sometimes obscurely marked, some-
times abrupt and conspicuous —into sap-wood and hear-wood.
The former is the popular name given to the outer and newer
layers of softer, more open, and bibulous wood. The early physi-
ologists named it alburnum from its white or pale color. Being
more or less sappy, or containing soluble organic matter, and
readily imbibing moisture, this part of the wood is liabie to decay,
and it is therefore discarded from timber used for construction.

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

INTERNAL STRUCTURE. 81

In all trees which have the distinction between the sap-wood and
heart-wood well marked, the latter acquires a deeper color, and
that peculiar to the species, such as the dark brown of the Black
Walnut, the blacker color of the Ebony, the purplish-red of Red
Cedar, and the bright yellow of the Barberry. These colors are
owing to special vegetable products, or sometimes to alterations
resulting from age. In the Red Cedar, the deep color belongs
chiefly to the medullary rays. In many of the softer woods, there
is little change in color of the heart-wood, except frem incipient
decay, as in the White Pine, Poplar, Tulip-tree, &c. The
heart-wood is no longer in any sense a living part: it may perish,
as it frequently does, without affecting the life or health of the
tree.

148. The Growth and Duration of the Bark, also the differences
in structure, are much more various than of the wood. Moreover,
the bark is necessarily subject to grave alterations with advanc-
ing age, on account of its external position; to distention from
the constantly increasing diameter of the stem within, and to
abrasion and decay from the influence of the elements without.
It is never entire, therefore, on the trunks of large trees; but
the dead exterior parts, no longer able to enlarge with the en-
larging wood, are gradually fissured and torn, and crack off in
strips or pieces, or disappear by slow decay. So that the bark
of old trunks bears only a small proportion in thickness to the
wood, even when it makes an equal amount of annual growth.

149. The three parts of the bark (142), for the most part
readily distinguishable in the bark of young shoots, grow inde-
pendently, each by the addition of new cells to its inner face, so
long as it grows at all. The green layer commonly does not
increase after the first year ; the opaque corky layer soon excludes
it from the light; and it gradually perishes, never to be renewed.
The corky layer usually increases for a few years only, by the
formation of new tabular cells : occasionally it takes a remarkable
development, forming the substance called Cork, as in the Cork
Vak, and the thin and parchment-like layers of the White and
Paper Birches.

150. The liber, or inner bark, continues its growth through-
out the life of the exogenous tree, by an annual addition from
the cambium-layer applied to its inner surface. Sometimes this
growth is plainly distinguishable into layers, corresponding with
or more numerous than the annual layers of the wood: often,
there is scarcely any trace of such layers to be discerned. In
composition and appearance, the liber varies greatly in different
plants, especially in trees and shrubs. That of Basswood or

82 MORPHOLOGY OF STEMS.

Linden, and of other plants with a similar fibrous bark, may be
taken as best representing the liber. Here it consists of alter-
nate strata of fibrous bast, and of the peculiar liber-cells called
sieve-cells, in which nourishing matter is especially contained
and elaborated. While the latter, or their equivalents, occur
and play an important part in all inner bark, the bast-cells are
altogether wanting in the bark of some plants, and are not pro-
duced after the first year in many others. The latter is the case
in Negundo, where abundant bast-cells, like those of Basswood,
compose the exterior portion of the first year’s liber, but none
whatever are formed in the subsequent layers. In Beeches and
Birches, also, a few bast-cells are produced the first year, but
none afterwards. In Maples, a few are formed in succeeding
years. In the Pear, bast-cells are annually formed, but in very
small quantity, compared with the parenchymatous part of the
liber. In Pines, at least in White Pines, the bark is nearly as
homogeneous as the wood, the whole liber, except what answers
to the medullary rays, consisting of one kind of cells, resembling
those of bast or of wood in form, but agreeing with the proper
liber-cells in their structure and markings.

151. The bark on old stems is constantly decaying or falling
away from the surface, without any injury to the tree; just as
the heart-wood within may equally decay without harm, except
by mechanically impairing the strength of the trunk. ‘There are
great differences as to the time and manner in which the older
bark of different shrubs and trees is thrown off. Some have
their trunks invested with the liber of 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 (Spirzea opulifolia), and of Grape-vines (except
Vitis vulpina), the liber lives only one season, and is detached
the following year, hanging loose in papery layers in the former
species, and in fibrous shreds in the latter.

152. While the newer layers of the wood abound in crude sap.
which they convey to the leaves, those of the inner bark abound
in elaborated sap, which they receive from the leaves and convey
to the cambium-layer or zone of growth. The proper juices and
peculiar products of plants are accordingly found in the foliage
and the bark, especially in the latter. Im the bark, therefore
(either of the stem or of the root), medicinal and other principles
are usually to be sought, rather thanin the wood. Nevertheless,
as the wood is kept in connection with the bark by the medullary
rays, many products which probably originate in the former are
deposited in the wood.

INTERNAL STRUCTURE. 83

153. The Living Parts of a tree or shrub, of the exogenous
kind, are obviously only these: 1st, The summit of the stem
and branches, with the buds which continue them upwards and
annually deveiop the foliage. 2d, The fresh roots and rootlets
annually developed at the opposite extremity. 35d. The newest
strata of wood and bark, and especially the interposed cambium-
layer, which, annually renewed, maintain a living communication
between the rootlets on the one hand and the buds and foliage
on the other, however distant they at length may be. These are
all that is concerned in the life and growth of the tree ; and these
are annually renewed. The branches of each year’s growth are,
therefore, kept in fresh communication, by means,of the newer
layers of wood, with the fresh rootlets, which are alone active in
absorbing the crude food of the plant from the soil. The fluid
they absorb is thus conveyed directly to the branches of the sea-
son, which develop leaves to digest it. And theSap they receive,
having been elaborated and converted into organic nourishing
matter, is partly expended in the upward growth of new branches,
and partly in the formation of a new laver of wood, reaching
from the highest leaves to the remotest rootlets.

154. Longevity of trees. As the exogenous tree, therefore,
annually renews its buds and leaves, its wood, bark, and roots,
— every thing, indeed, that is concerned in its life and growth, —
there seems to be no necessary cause, inherent in the tree itself,
why it may not live indefinitely. Some trees are known to have
lived for one and two thousand years, and some are possibly
older.1 Equally long may survive such endogenous trees as the

Dragon tree (Draczena), which have provision for indefinite in-
crease in diameter (138), and for the production of branches.
The famous Dragon tree of Orotava, in Teneriffe, now destroyed
by hurricanes and other accidents, had probably reached the age
of more than two thousand years.

155. On the other hand, increase in height, spread of branches
and length of root, and extension of the surface over which the
annual layer is spread, are attended with inevitable disadvantage,
which must in time terminate the existence of the tree in a way
quite analogous to the death of aged individual animals, which
is not directly from old age, but from casualties or attacks to

1 The subject of the longevity of trees has been discussed by DeCandolle,
in the “ Bibliotheque Universelle” of Geneva, for May, 1831, and in the second
volume of his “ Physiologie Végétale ;” more recently, by Alphonse DeCan-
dolle in the “ Bibliotheque 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.

8+ MORPHOLOGY OF STEMS.

which the aged are either increasingly incident or less able to
resist. A tree like the Banyan (59, Fig. 71), which by aerial
roots continues to form new trunks for the support and sustenance
of the spreading branches, and thus ever advances into new soil,
has a truly indefinite existence ; but, then, it becomes a forest,
or is to be likened to a colony propagated and indefinitely in-
creased by suckers, offsets, or other subterranean shoots. So
the question of the secular continuation of the individual plant
becomes merged in that of continuation of the race, — at least of
a bud-propagated race, — the answer to which is wholly in the
domain of conjecture.’ However this may be, it is evident that
a vegetable of the higher grade is not justly to be compared with
an animal of*higher grade; that individuality is incompletely
realized in the vegetable kingdom ;? that rather

156. The Plant is a Composite Being, or Community, lasting, in
the case of a treg, through an indefinite and often immense num-
ber of generations. These are successively prodaced, enjoy a
term of existence, and perish in their turn. Life passes onward
continually from the older to the newer parts, and death follows,
with equal step, at a narrow interval. No portion of the tree is
now living that was alive a few years ago; the leaves die annu-
ally and are cast off, while the mternodes or joints of stem that
bore them, as to their wood at least, buried deep in the trunk
under the wood of succeeding generations, are converted into
lifeless heart-wood, or perchance decayed, and the bark that
belonged to them is thrown off from the surface. It is the aggre-
gate, the blended mass alone, that long survives. Plants of
single cells, and of a definite form, alone exhibit complete indi-
viduality ; and their existence is extremely brief. The more
complex vegetable of a higher grade is not to be compared with
the animal of the highest organization, where the offspring always
separates from the parent, and the individual is simple and indi-
visible. But it is truly similar to the branching or arborescent
coral, or to other compound animals of the lowest grade, where
successive generations, though capable of living independently
and sometimes separating spontaneously, yet are usually devel-
oped in connection, blended in a general body, and nourished
more or less in common. ‘Thus, the coral structure is built up
by the combined labors of a vast number of individuals, — by
the successive labors of many generations. The surface or the
recent shoots only are alive; beneath are only the dead remains

1 See Darwiniana, xii. 338-355.
2 As, perhaps, was first explicitly stated by Engelmann, in his inaugural
essay, De Antholysi Prodromus, Introduction, § 4.

MORPHOLOGY OF LEAVES. 85

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

Section IV. Or Leaves.
§ 1. THerr NATURE AND OFFICE.

157. Leaf (Lat. Yolium, in Greek form Phyllum), as a botani-
cal term, has on the one hand a comprehensive, on the other a
restricted sense. In its commonest sense, as used in descriptive
botany, it denotes the green blade only. Yet it is perfectly
understood that the footstalk is a part of the leaf, and therefore
that the phrase ‘* leaves cordate,” or the like, is a short way of
saying that the blade of the leaf is cordate or heart-shaped.
Moreover, two appendages, one on each side of the base of the
footstalk, when there is any, are of so common occurrence that
they are ranked as a proper part of the organ. So that, to the
botanist, a typical leaf consists of three parts: 1, BLApE or
Lamia; 2, Foor-statk or Lear-sraLk, technically PErio.e ;
3, A pair of Stieutes. (Fig. 142.)

158. The blade, being the most important part of an ordinary
leaf, may naturally be spoken of as the whole. Petiole and
stipules are indeed subsidiary when present, and are not rarely
wanting. Yet sometimes they usurp the whole function of foli-
age, and sometimes there is no such distinction of parts.

159. Physiologically, leaves are green expansions borne by the
stem, outspread in the air and light, in which assimilation (3)
and the processes connected with it are carried on. Vegetable
assimilation, — the most essential function of plants, being the
conversion of inorganic into organic matter, —takes place in
all ordinary vegetation only in green parts, and in these when
exposed to the light of the sun. And foliage is an adaptation
for largely increasing the green surface. But stems, when green,
take part in this office in proportion to the amount of surface,
sometimes monopolize it, and in various cases increase their
means of doing so by assuming leaf-like forms. (126-129.)
Leaves, especially in such cases, may lose this function, appear
only as useless vestiges, or may be subservient to various wholly

86 MORPHOLOGY OF LEAVES.

different uses. Form and function, therefore, are not sure indi-
cations of the true nature of organs.

160. Morphologically, and in the most comprehensi¥e sense,
leaves are special lateral outgrowths from the stem, definitely
and symmetrically arranged upon it; in ordinary vegetation and
in the most general form constituting the assimilating apparatus
(or foliage), but also occurring in other forms and subserving
various uses. Sometimes these uses are combined with or sub-
sidiary to the general function of foliage ; sometimes the leaf is
adapted to special uses only. So the botanist — recognizing the
essential identity of organs, whatever their form, which appear
in the position and conform to the arrangement of leaves —
discerns the leaf in the cotyledons of a bean or acorn, the scale of
a lily-bulb or the coat of an onion, the scale of a winter bud, and
the petal of a blossom. ‘Therefore, while expanded green leaves
(which may be tautologically termed jfoliage-leaves) are taken as
the proper type, the common name of leaves, in the lack of any
available generic word, is in morphological language extended to
these special forms whenever it becomes needful to express their
phylline or foliar nature.

161. In the morphological view, all the plant’s organs except-
ing roots (and excepting mere superficial productions, such as
hairs, prickles, &c.), belong either to stem or to leaves, are
either cauline or phylline in nature. ‘To the latter belong all the
primary outgrowths from nodes, all lateral productions which
are not axillary... Whatever is produced in the axil of a leaf is
cauline, and when developed is a branch.

162. The Duration of Leaves is transient, compared with that
of the stem. They may be fugacious, when they fall off soon
after their appearance; deciduous, when they last only for a
single season; and persistent, when they remain through the cold
season, or other interval* during which vegetation is interrupted,
and until after the appearance of new leaves, so that the stem is
never leafless, as in Evergreens. In many evergreens, the leaves
have only an annual duration; the old leaves falling soon after
those of the ensuing season are expanded, or, if they remain
longer, ceasing to bear any active part in the economy of the
vegetable, and soon losing their vitality altogether. In Pines
and Firs, however, although there is an annual fall of leaves
either in autumn or spring, yet these were the produce of some

1 There are cases in which this rule is of difficult application, or is seem-
ingly violated, sometimes by the suppression of the subtending leaf, as in the
inflorescence of Crucifere, rarely in other ways, to be explained in the
proper places.

none

THEIR STRUCTURE AND FORMS. 87

season earlier than the last; and the branches are continually
clothed with the foliage of from two to five, or even ten or more
successive years. On the other hand, it is seldom that all the
leaves of an herb endure through the whole growing season, the
earlier foliage near the base of the stem perishing while fresh
leaves are still appearing above. In our deciduous trees and
shrubs, however, the leaves of the season are mostly developed
within a short period, and they all perish in autumn nearly
simultaneously.

163. Leaves soon complete their growth, and have no power
of further increase. Being organs for transpiration, a very large
part of the water imbibed by the roots is given out by the foliage,
leaving dissolved earthy matters behind. Assimilation can take
place only in fresh and vitally active tissue. It is incident to all
this that leaves should be of only transient duration, at least in
their active condition.

164. Defoliation. The leaves of most Dicotyledons and some
Monocotyledons separate from the stem and fall by means of an
articulation at the junction with the stem, which begins to form
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, such as is seen
in Fig. 81, 85, 91. Some leaves, notably those of Palms,
Yucca, and other endogens, die and wither on the stem, or wear
away without falling.

165. In temperate climates, defoliation mostly takes place at
the approach of winter. In warmer climates having only winter
rain, this occurs in the hot and dry season.

166. Normal Direction or Position. The leaf-blade is expanded
horizontally, that is, has an upper and an under surface. When
erect, the upper surface faces the axis which bears it. To this,
there are many seeming but no real exceptions; that is, none
which are not explicable as deviations or changes from the normal
condition. (215-217.)

§ 2. THerr Structure AND Forms AS ORGANS OF ASSIMILATION
orn VEGETATION, @. e. AS FOLIAGE.

167. The Internal Structure or Anatomy of the leaf needs here
to be examined so far as respects its obvious parts and their
general composition. The leaf, like the stem, is composed of
two elements (131), the cellular and the woody. The cellular
portion is the green pulp or parenchyma, and in this the work
of assimilation is carried on. The woody is the fibrous frame-

85 MORPHOLOGY OF LEAVES.

work, the separate parts or ramifications of which form what are
variously called the 77s, — a sufticiently proper term, — nerves or
veins. ‘The latter names may suggest false analogies ; but they
are of the commonest use in descriptive botany. That of veins,
and of its diminutive, vein/ets, for the smaller ramifications, is
not amiss; for the fibrous framework not only gives firmness
and support to the softer cellular apparatus, 7. e. forms ribs,
but serves in the leaf, as it does in the stem, for the more rapid
conveyance and distribution of the sap. The subdivisions con-
tinue beyond the limits of unassisted vision, until the fibro-vas-
cular bundles are reduced to attenuated fibres ramified through
the parenchyma. In leafstalks, the woody bundles are parallel,
not ramified, and arranged in various ways; in Exogens usually
so as to form in cross-section an arc or an incomplete or com-
plete ring. In leaves serving as foliage or organs of assimilation,
the blade is the important part, and this only is here regarded.

168. The characteristic contents of these cells of parenchyma
are grains of chlorophyll (142°), literally leaf-green, to which
the green color of foliage is wholly owing, and which may be
regarded as the most important of all vegetable products ;
because it is in them (or in this green matter, whatever its form,
189) that all ordinary assimilation takes place. As it acts only
under the influence of light, the expanded leaf-blade may be
viewed as an arrangement for exposing the largest practicable
amount of this green matter — the essential element of vegeta-
tion — to the light and air.

169. The Parenchyma-cells, constituting the green pulp, are
as eal Gas can | | themselves arranged in accordance

Eee yseepees With this adaptation. The upper stra-
tum is mostly of oblong cells, compactly
arranged in one or more layers, their
longer diameter perpendicular to the
surface. The stratum next the lower
surface of the leaf consists of loosely
arranged cells, with longer diameter
usually parallel to the plane of the leaf,
often irregular in form, and so disposed
as to leave intervening sinuous air-spaces freely permeating all

FIG. 136. A magnified section through the thickness of a leaf of Iicium Flori-
danum, showing the irregular spaces or passages between the cells, which are small in
the upper layer of the green pulp, the cells of which (placed vertically) are well com-
pacted, so as to leave only minute vacuities at their rounded ends; but the spaces are
large and copious in the rest of the leaf, where the cells are very loosely arranged:
also the epidermis or skin of the upper (a) and of the lower surface of the leaf (0d),
composed of perfectly combined and thick-walled empty cells,

THEIR. STRUCTURE AND FORMS. 89

that part of the leaf. (Fig. 136, 137.) Hence in good part the
deeper green hue of the upper, and the paler of the lower face
of leaves.

170. Epidermis. The whole surface of leaves, as of young
stems, is invested with a translucent membrane, composed of
one or sometimes two or three layers of empty and rather-
thick-walled cells. This is the skin or epidermis, which is so
readily separable from the succulent tissue of such leaves as
those of Stonecrop and other species of Sedum. It is of a single
layer in the Illicium (Fig. 136) and Lily (Fig. 137); of as
many as three in the firm leaf of the Oleander; is generally

187

hard and thick in such coriaceous leaves as those of Pittosporum
and Laurustinus, which thereby the better endure the dry air of
rooms in winter.

171. Stomata or Breathing-pores.t| The epidermis forms a
continuous protective investment of the leaf except where certain
organized openings occur, the stomata. They are formed by a
transformation of some of the cells of the epidermis ; and consist
usually of a pair of cells (called guardian-cells) , with an opening
between them, which communicates with an air-chamber within,
and thence with the irregular intercellular spaces which permeate
the interior of the leaf. Through the stomata, when open, free
interchange may take place between the external air and that

1 The technical name has been anglicized stomates, singular stomate, which
has no advantage over the proper Greek, sing. sfoma, pl. sfomata.
FIG. 137. A magnified section through the thickness of a minute piece of the leaf

of the White Lily of the gardens, showing also a portion of the uuder side with some
breathing-pores, stomata.

90 MORPHOLOGY OF LEAVES.

within the leaf, and thus transpiration be much facilitated.
When closed, this interchange will be interrupted or impeded.
The mechanism of stomata is somewhat
recondite, and will be illustrated in the
anatomical and physiological volume of
this series.

172. It is only when leaves assume
a vertical or edgewise position that the
stomata are in equal numbers on both
faces of a leaf. Ordinarily. they occupy
or most abound on the lower face, which
is turned away from the sun; but in certain coniferous trees the
reverse of this is true. In the Water Lilies (Nymphzea, Nuphar),
and other leaves which float
upon the water, the stomata all
belong to the upper surface.
Leaves which live under water,
where there can be no evapora-

a a va tion, are destitute, not only of
stomata, but usually of a distinct epidermis also. The number
of the stomata varies from 800 to about 170,000 on the square
inch of surface in different leaves. In the Apple, there are said
to be about 24,000 to the square inch (which is under the average
number, as given in a table of 36 species by Lindley) ; so that
each leaf of that tree would present about 100,000 of these
orifices. The leaf of Dragon Arum is said to have 8,000
stomata to a square inch of the upper surface, and twice that
number in the same space of the lower. That of the Coltsfoot
has 12,000 stomata to a square inch of the lower epidermis,
and only 1.200 in the upper. That of the White Lily has
from 20,000 to 60,000 to the square inch on the lower sur-
face, and perhaps 3,000 on the upper; and they are so re-
markably large that they may be discerned by a simple lens of
an inch focus.

172. Venation, the veining of leaves, &c., relates to the mode
in which the woody tissue, in the form of ribs, veins, &e., is
distributed in the cellular. There are two principal modes, the
parallel-veined and the reticulated or netted-veined. The former
is especially characteristic of plants with endogenous stem and
monocotyledonous embryo, and also of gymnospermous trees,

FIG. 138. A highly magnified piece of the epidermis of the Garden Dalsam, with
three stomata (after Brongniart).

FIG. 139. Magnified view of the 10,000th part of a square inch of the epidermis of
the lower surface of the leaf of the White Lily, with its stomata. 140. A single stoma,
more magnified. 141. Another stoma, widely open.

i i eel

THEIR STRUCTURE AND FORMS. 91

which have exogenous stems and at least dicotyledonous
embryos. The latter prevails in ordinary plants with exogenous
stem and dicotyledonous embryo.

173. Parallel-veined or Neryed leaves (of which Fig. 143 is
an illustration) have a framework of simple ribs (called by the
earlier botanists zerves, a name still used in descriptions), which
run from the base to tip, or sometimes from a central strong rib
to margin of the leaf, in a generally parallel and undivided way,
and sending off or connected by minute veinlets only. Grasses,

142 143

Lily of the Valley, and the like, illustrate the commoner mode
in which the threads of wood run from base to apex. The
Banana and Canna are familiar illustrations of a mode not un-
common in tropical or subtropical endogens, in which the threads
or ‘‘nerves” run from a central rib (midrib) to the margin.
Parallel-veined leaves are generally entire, or at least their
margins not toothed or indented. The principal exception to
this occurs when the ribs or the stronger ones are few in number
and radiately divergent, as in the flabelliform leaves of Fan-
palms, a peculiar modification of the parallel-veined type.
Between leaves with nerves wholly of basal origin, and those
with nerves all springing from a midrib, there are various grada-
tions, and also in respect to curving. But parallel-veined or
nerved leaves may be classified into

FIG. 142. A leaf of the Quince, of the netted-veined or reticulated sort, with
blade (4), petiole or leaf-stallk (), and stipules (sf).
FIG. 143. Parallel-veined leaf of the Lily of the Valley, Convallaria majalis.

92 MORPHOLOGY OF LEAVES.

Pasal-nerved, that is, with the nerves all springing from the
base of the leaf, and

Costal-nerved, springing from a midrib or costa. Either may be

Rectinerved, the nerves running straight from origin to apex
or margin of the leaf, as the case may be ;

Curvinerved, when curving in their course, as in the leaves
of Funkia and in Canna ;

Flabellinerved, where straight nerves and ribs radiate from the
apex of the petiole, as in Fan-palms and the Gingko tree.

174. In typical parallel-veined leaves, all reticulation is con-

fined to minute and straight cross-veinlets: in many, these are.

coarser, branching, and reticulated ; in some, as in Smilax and
Dioscorea, only the primary ribs or strongest nerves are on the
parallel-veined plan ; the space between being filled with reticu-
lations of various strength; thus passing by gradations into

175. Reticulated or Netted-veined leaves. In most of these,
from one to several primary portions of the framework are
particularly robust, and give origin to much more slender ram-
ifications, these to other still smaller ones, and so on. The
strong primary portions are Ries (cosfe) ; the leading ramifi-
cations, VErNs (vere) ; the smaller and the ultimate subdivisions,
VEINLETS (venul@). All or some of the veins and veinlets are
said to anastomose, 7. e. variously to connect with those from
other trunks or ribs, apparently in the manner of the veins and
arteries of animals, forming meshes. But, as there is no opening
of calibre of one into another, the word is etymologically rather
misleading. More properly, it is said that the veins or veinlets
form reticulations or net-work. A primary division of reticulated
leaves, and indeed of nerved leaves also, into two classes, is
founded upon the number of primary ribs.

176. There may be only a single primary rib; this traversing
the blade from base to tip through its centre or axis (as in Fig.
142, 152-156) is called the Mrpris. There may be others, gen-
erally few (one, two, three, or rarely four), rising from the apex
of the petiole on each side of the midrib, running somewhat par-
allel with it or more or less diverging from it: these are lateral
ribs. Among parallel-veined leaves, the Banana, Canna, &c.,
have a single rib, from which the veins (in the older nomencla-
ture here called nerves) all proceed. Most Lilies and the like
have several approximately parallel ribs, but the midrib pre-
dominant: in other cases, the midrib is no stronger than the
others. In Fan-palms, the ribs are radiately divergent, giving
a fan-shaped or rounded outline to the blade. In reticulated
leaves, in which the veins all spring from the ribs, the two

= es

THEIR STRUCTURE AND FORMS. 93

classes into which they divide are the pinnately veined and the
palmately veined.

177. Pinnately or Feather-veined (or Penninerved) leaves
are accordingly those of which the veins and their subdivisions
are side branches of a*ingle central rib (midrib), which traverses
the blade from base to apex; the veins thus being disposed in
the manner of the plume on the shaft of a feather. (Fig. 142,
152, &e.) Sometimes these continue straight and undiminished
from midrib to margins (straight-veined, as in Beech and Chest-
nut, Fig. 152), sending off only small lateral veinlets ; some-
times they ramify in their course into secondary or tertiary veins,
and these into veinlets. Pinnate venation in reticulated leaves
naturally belongs to leaves which are decidedly longer than wide.

178. Some of the primary veins, commonly among the lower,
may be stronger than the rest, and thus take on the character of
ribs, or by gradations pass into such. The leaf of the common
annual Sunflower (Fig. 155) becomes in this way ¢riple-ribbed or
tripli-nerved. ‘The appearance of a second pair of such strength
ened veins makes the venation guintupli-ribbed ov quintupli-nerved.

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

179. Palmately, Digitately, or Radiately Veined (or Palmi-
nerved) class, of which leaves of common Maples and the Vine
are familiar examples. (Also Fig. 158-160, &c.) In these

FIG. 144-157. Various forms of simple leaves, explained in the text and in the
Glossary.

94 MORPHOLOGY OF LEAVES.

there are three, five, seven, or sometimes more ribs of equal
strength, the central being the midrib, and each with its system
of veins which ramify and form meshes in the interspaces. Here
the whole woody portion of the leaf divides equally into a num-
ber of parts upon leaving the petiole or entering the blade. The
ribs there commonly diverge more or less in a palmate or digitate
manner (7. e. like outspread fingers of the hand, or the claws of a
bird, or like radii of more or less of a circle) : so, in the corre-
lation df outline and venation, this class of veining goes with

roundish circumscription. This is not so true, however, in a
special case, viz., where the ribs, however divergent below, curve
forward and all run to the apex of the blade, thus imitating the
parallel-veined system, as in Rhexia and generally in the family
of which that genus is the single northern representative.’

180. Forms as to Outline, &e. DeCandolle conceived the shape
of leaves (both the general circumscription and the special con-
figuration) to depend on the distribution of the ribs and veins,
and quantity of the parenchyma in which these were outspread,
—-a too mechanical view, and not conformable to the history of
development. This proves that the framework is adapted to the
parenchyma, which grows and shapes the organ in its own way,
rather than the parenchyma to it. It were better to say that the

1 In Linnean terminology, palmate and digitate referred to particular out-
line only, and were separately used to denote extent of division, — palmate, not
divided down to the petiole, digitate, when divided, like the claws of a bird,
quite down the base. DeCandolle generalized the use of the former term,
and ever since the two have been used interchangeably.

FIG. 158-166. Various forms of simple, chiefly palmately veined leaves.

THEIR STRUCTURE AND FORMS. 95

two elements of the structure are correlated. Descriptive terms
applied. to leaves are equally applicable to all expanded organs
or parts, and indeed to all outlines. Some leading forms are
here enumerated ; and all are defined in the Glossary.

181. As to general Circumscription, proceeding from narrower
to broader shapes, and then to those with either narrowed or
notched base, leaf-blades are

167 168

170

172

Linear, when narrow, several times longer than wide, and of
about the same breadth throughout. (Fig. 167.)

Lanceolate, or Lance-shaped, when several times longer than
wide, and tapering upwards (Fig. 153, 168), or tapering both
upward and downward.

Oblong, when nearly twice or thrice as long as broad. (Fig. 169.)

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

Oval, the same as broadly elliptical, or elliptical with the

breadth considerably more than half the length.
Ovate, when the outline is like a section of a hen’s-egg length-
wise, the broader end being downward. (Fig. 171, 155.)
Obovate, inversely ovate, or
ovate with the narrower end
toward the base, the broader
Cuneate, or Cuneiform, that is, We a eed i oad cee and
tapering by straight lines to an acute base. (Fig. 176, 148.)
Spatulate, rounded above, long and narrow below, like a
Oblanceolate, inverted lance-shaped, 7. e. such a lanceolate
leaf as that of Fig. 168, but with the more tapering end at base,
as in Fig. 173. To those who restrict the term lanceolate to the

Orbicular, or Rotund, circu-
lar in outline, or nearly so.
(Fig. 160.)
upward. (Fig. 175, 145.) eee oe
spatula. (Fig. 174, 147.)
sense of a narrow leaf tapering equally in both directions, the

FIG. 167-176. Outlines of various simple leayes.

G6 MORPHOLOGY OF LEAVES.

term oblanceolate is superfluous. The following terms desig-
nate leaves with a notched instead of narrowed base.

Cordate, or Heart-shaped, when a leaf of an ovate form, or
something like it, has the outline of its rounded base turned in
(forming a notch or sinus) where the stalk is attached, as in
Fig. 172, 151. Also Fig. 149, Pontederia, a leaf of the parallel-
veined class.

Reniform, or Kidney-shaped, like the last, only rounder and
broader than long. (Fig. 158.)

Auriculate, or Eared, having a pair of small and blunt pro-
jections, or ears, at the base, as in Magnolia Fraseri, Fig. 178.

Sagittate, or Arrow-shaped, where such ears are acute and turned
downwards, while the
main body of the blade
tapers upwards to a
point, as in the com-
mon species of Sagit-
taria or Arrow-head,
and in the Arrow-
leaved Polygonum.
(Fig. 165, 177.)

flastate, or Halberd-
shaped, when such
lobes at the base point outwards, giving the leaf the shape of the
halberd of the olden time, as in Polygonum arifolium (Fig. 179)
and Sorrel, Fig. 163.

182. Peltate or Shield-shaped leaves are those in which a blade
of rounded or sometimes of other shape is attached to the petiole
by some part of the lower surface, instead of the basal margin:
those of Water-shield or Brasenia, of Nelumbium, and of Hydro-
cotyle umbellata are marked examples. The anomaly is mor-
phologically explained by a comparison with deeply cordate or
reniform leaves haying a narrow sinus, such as those of Nym-
pheea or Water Lily, and by supposing a union of the approxi-
mated edges of the sinus. Fig. 159 and 160, from two species
of Hydrocotyle, one with open and the other with closed sinus
obliterated by the union, illustrate this.

185. As to Extremity, whether base or apex, there are several
descriptive terms, expressive of the principal modifications ;
such as

Acuminate, tapering, either gradually or abruptly, into a
narrow more or less prolonged termination. (Fig. 180.)

FIG. 177-179. Sagittate, auriculate, and hastate leaves.

i ee ee

THEIR STRUCTURE AND FORMS. aK

Arute, ending in an acute angle, without special tapering, as
in Fig. 181.

Obtuse, ending with a blunt or roundish extremity, Fig. 182.

Truncate, with termination as if cut off by a straight transverse
line, as in Fig. 183.

Retuse, with an obtuse extremity slightly depressed or re-enter-
ing, as in Fig. 184.

Emarginate, with a more decided terminal notch, Fig. 185.

Obcordate, inversely heart-shaped, 7. e. like cordate, but the
broader end and its strong notch at apex instead of base, Fig.
186. This and the following terms are applicable to apex only.

Mucronate, abruptly tipped with a small and short point, like a
projection of the midrib, as in Fig. 188.

Cuspidate, tipped with a sharp and rigid point, as in Fig. 187.

184. As to Margin or special Outline, the terminology proceeds
upon the convenient supposition of a blade with quite entire margin,
but subject to incisions, which give rise to notches or clefts, if we
regard the sinuses ; or to teeth, lobes, segments, &c., if we regard
the salient portions between the sinuses. The ribs, or the stronger
veins, &c., commonly terminate in the teeth or lobes ; but in Cicuta
maculata, and in a few other cases, they run to the notches.

185. Dentation relates to mere marginal incision, not extend-
ing deeply into the blade. The blade is said to be

Entire} when
the margin is com-
pletely filled out
to an even line, as
in Fig. 173-179.

Serrate, when
with small and |
sharp teeth direct-} >}
ed forward, a
the teeth ofa saw, \
as in Fig. 189. Se

Serrulate is the ‘159 <8 191
diminutive of serrate, and is eaten to minutely serrate.

1 Integerrimus-a-um, or quite entire, is the term in Latin terminology. Inte
ger Means undivided or not lobed.
FIG, 180-188. Terminations of leaves.
FIG. 189-194. Dentation of pinnately veined leaves.

7

98 MORPHOLOGY OF LEAVES.

Dentate, or Toothed, a general term for toothing, specially
applied to the case of salient teeth which are not directed for-
ward or towards the apex of the blade, Fig. 190.

Crenate, or Scalloped, the same as dentate or serrate, but with
teeth much rounded, Fig. 191.

Repand, or Undulate, when the margin is a wavy line, bending
slightly inward and outward, Fig. 192.

Sinuate, when this wavy line is stronger or distinctly sinuous,
as in Fig. 193.

Incised, when cut by sharp and irregular incisions more or less
deeply, Fig. 194. This is intermediate between dentation and

186. Lobation or Segmentation. When the blade is more
deeply penetrated by incisions from the margin, that is, when
the spaces between the ribs or principal veins are not filled to
near the general outline, it is said to be Jobed, cleft, parted, or
divided, according to the degree of separation ; and the portions
are called lobes, segments, divisions, &e. The most general name
for such parts of any simple blade is that of /vbes. More par-
ticularly a leaf-blade, or other body, is said to be

Lobed, when the division extends not more than half way down,
and either the sinuses or the lobes are rounded ;

Cleft, when the division is half way down or more, and the
lobes or sinuses narrow or acute ;

Parted, when the divisions reach almost, but not quite, to the
base or the midrib ;

Divided, when they sever the blade into distinct parts, which
makes the leaf compound. (193.)

i187. Lope is the common name of one of the parts of a simple
blade, especially when there is only one order of incision. But

when there are more, as when a leaf is divided or parted and—

these primary lobes again lobed or cleft, the lobes of first order
are commonly called SeGMENTs (sometimes divisions or partitions),
and the parts of these, Lobes. Or the lobes may be designated
as primary, secondary, tertiary, &c. Ultimate portions or small
lobes may be called Lobules or Lobelets. Also the portions of a
quite divided blade take the name of Leaflets. By proper selec-
tion of terms, the degree of division or lobing may thus be
expressed in a single word. :

188. As to Number of parts, this may be tersely expressed by
combination with the adjective term applicable to the degree ; as,
Two-lobed, Three-lobed, Five-lobed, Many-lobed, &c. ; or Two—Five-
cleft, Many-cleft, &c., in Latin form Bifid, Trifid, Multifid, &e. ;
Two-five-parted, &c., according to the number of divisions
which extend almost to the base or axis; Two—Five-divided (in

—————

THEIR STRUCTURE AND FORMS, 99

Latin form Bisected, Trisected, &c.), when there are two or three
or more complete divisions of the blade.

189. As to Arrangement of parts, this may be simply and best
expressed by taking into account the nature of the venation or
the distribution of the ribs, &e., which controls or is co-ordinated
with the disposition of the lobes. Pinnately veined leaves,
when lobed, must needs have the incisions directed to the mid-
rib; palmately-veined or radiated, to the apex of the petiole.
The lobes or divisions of the first will be pinnately, of the second

palmately disposed. Accordingly, the three leaves of as many
species of Oak, Fig. 195, 196, and 197, represent respectively a
pinnately lobed, pinnately cleft, and pinnately parted leaf, while the
accompanying leaf of Celandine, Fig. 198, is pinnately divided.
The first three, however, when the degree of incision 1s not par-
ticularly in question, usually pass under the common term of
pinnatifid, Fig. 195 moderately, Fig. 197 deeply. The number
of lobes, when definitely marked, may come into the descriptive
phrase, as pinnately 7-lobed, pinnately 7-cleft, parted, or divided,
as the case may be.

190. Similarly, Figures 199 to 202 represent, respectively, a
palmately three-lobed, three-cleft, three-parted, and three-divided, or,
in Latin form, trilobate, trifid, tripertite, and trisect or trisected
leaf. Fig. 166 is a palmately 5-parted leaf; Fig. 164, palmately

FIG. 195-198. Pinnately lobed, cleft, parted, and divided leaves.
FIG, 199-202. Palmately 3-lobed, cleft, parted, and divided leaves.

100 MORPHOLOGY OF LEAVES.

multifid, &c. Fig. 162, a leaf of Dragon Arum, is palmately
9-parted. But, as the lateral sinuses are not so deep as the
others, the leaf is said to be pedately parted, or pedate, in the
early terminology.

191. Moreover, as the lobes or divisions of a leaf may be
again similarly lobed or parted, &c., this composition may be
indicated by the prefix twice, thrice, &¢., as twice pinnatifid or
bipinnatifid, thrice pinnately parted, thrice palmately parted, and
the like. Thus, a word or two, or a short phrase, may describe
even a complex leaf, so as to convey a perfectly clear and defi-
nite idea of its conformation.

192. A distinction should now be drawn between simple and
compound leaves. The distinction cannot be both natural and
absolute ; for the one may pass variously into the other. Simple
leaves, which have been thus far considered, have a single lamina
or blade, which may, however, at one extreme be entire, at the
other many-parted, and even several times divided.

193. Compound Leaves are those which have from two to
many distinct blades, on a common leafstalk. These blades,
called LearLets, may be sessile on the common leafstalk, or they
may have leafstalks of theirown. As the leaf very commonly
separates in age by an articulation of its petiole with the stem,
so leaflets are commonly more or less articulated with the com-
mon petiole. When the leaf, with its petiole, falls from the
stem, the leaflets may as completely separate from the common
petiole. They do not always do this. Divided leaves, such as
those of Fig. 198 and 202, though ranked among the simple
sorts, are compound in the sense of having distinct blades,
but without articulation. Some of these blades are apt to be
confluent ; that is, a divided leaf is often in part merely parted,
as in the upper portion of Fig. 198. Such leaves are so inter-
mediate between simple and compound that it becomes indiffer-
ent, or a matter of convenience to be settled by analogy, under
which head or by what language they shall be described. How-
ever, most leaves are so constituted as to leave no doubt whether
they are simple or compound.

194. The leaflets of a compound leaf being homologous with
the lobes or segments of a simple leaf, indeed being such segments
fully isolated, the two sorts fall under the same types. A _pin-
nately veined simple leaf is the homologue of one kind of com-
pound leaf; a radiately veined leaf, of the other. That is,
compound leaves are either pinnate or palmate.

195. Pinnate Leaves (Fig. 205-205) are those in which the
leaflets are arranged along the sides of a petiole, or rather of its

THEIR STRUCTURE AND FORMS. 101

prolongation, the Ruacuts, which answers to the midrib of a pin-
nately veined simple leaf. There are three principal sorts, and
some subordinate ones. That is, a pinnate leaf may be

Impari-pinnate, or pinnate with an odd leaflet, 7. e. a terminal
one, as in Fig. 203; and this is the commoner case.

Cirrhiferous Pinnate, or pinnate with a tendril (Fig. 204), as
in the proper Pea tribe and Bignonia. Here either the termi-
nal leaflet only, or the upper lateral leaflets also, are replaced
by tendrils.

Pari-pinnate, or Abruptly Pinnate, destitute of a terminal leaflet
or of any thing answering to it, as in Fig. 205.

Interruptedly Pinnate denotes merely a striking inequality of
size among the leaflets : Lyrately Pinnate, one in which the termi-
nal leaflet is largest and the lower small.

196. Palmate or Digitate Leaves (Fig. 206, 93) are those in
which the leaflets all stand on the
summit of the petiole. Digitate
(fingered) was the old name, when
the term palmate was restricted to
a simple but palmately lobed leaf
of this type. But since the time
of DeCandolle the two names have
been used interchangeably. Pal-
mate leaves have no primary dis-
tinction into sorts, except as to
the number of leaflets. These can
never be very numerous; but there are fully a dozen in some

FIG. 203. An impari-pinnate or odd pinnate leaf. 204. Pinnate with a tendril
205. Abruptly pinnate leaf of a Cassia.
FIG. 206. Palmately or digitately 5-foliolate leaf of a Buckeye, Zsculus.

102 MORPHOLOGY OF LEAVES.

Lupines. More commonly there are only five to nine, or only
three, rarely two, or even a single one.

197. Number of leaflets may be indicated by an adjective
expression composed of the proper Latin numeral prefixed to
foliolate (Foliolum, diminutive of foliwm, answering to leaflet).
Thus, bifoliolate, of two leaflets ; trifoliolate, of three leaflets;
guadrifoliolate, of four; quinquefoliolate, of five: plurifoliolate, or
multifoliolate, of several or numerous leaflets, &c. These terms
are still more descriptive when accompanied by the word pin-
nately or palmately, indicative of the kind of compound leaf; as,
palmately or digitately trifoliolate (common Clover-leaf, Fig. 211),
or 5-foliolate, as in Buckeye (Fig. 206), and so on. Also, pinnately
16-folvolate, as in Fig. 205, or 17%-foliolate, as in Fig. 203;
pinnately trifoliolate,as in Phaseolus, and in the low Hop-Clover,
Trifolium procumbens.'

198. But, in either class of compound leaves, the leaflets may
be reduced to a minimum number. A pinnately trifoliolate leaf
is one of the impari-pinnate kind reduced to three leaflets, to one
pair and the odd one; and this is distinguished from a palmately
trifoliolate leaf by the attachment of the pair at some distance
below the apex of the petiole, and by the articulation above this,
which marks the junction of the terminal leaflet’s petiole (or its
base, if sessile) with the rhachis or common petiole.

199. Unifoliolate compound leaves (by no means a direct con-
tradiction in terms) are by this articulation distin-
guished from simple leaves which they simulate.
See the leaf of the common Barberry, Fig. 207.
In other species, of the Mahonia section, the leaves
are all pinnately 5-9-foliolate, with well-developed
common petiole: in the true Berberis, they are all
thus reduced to the terminal and long-petiolulate
leaflet, on an almost obsolete petiole. Orange and
Lemon leaves are in similar case, but with the joint
close to the blade. A comparison with near rela-
tives shows that these are also unifoliolate leaves
of the pinnate kind; though this could not be ascertained by
inspection.

200. Decompound or Twice and Thrice Compound Leaves. These
are to once pinnate or once palmate leaves what the latter are to

1 In pinnate leaves, each leaflet usually has its opposite fellow, and the
number may be indicated by the pairs, as unijugate, bijugate, trijugate, and
plurijugate, according to the number of juga, or pairs.

FIG. 207. Unifoliolate leaf of Berberis vulgaris, with partial petiole articulated to
the extremely short true petiole.

THEIR STRUCTURE AND FORMS. 103

simple leaves. As leaflets may be toothed, lobed, or parted, so
what answers to a single leaflet may appear as leaflets of a second,
or again of a third, or even of a fourth order. Decompound is a
good general name for all more than once compounded leaves ;

Ww VY E&
° / ) J bes ry \\ \ vv 7
\ f (\\ \\ =

\ NN 77s Ze)
; SN /// SX /, WA
S aN
QY Yj XS i, a ee =>
WYO SESE (VS
A — =. fe WJ oD
SVG SW ea
SI Lp VOLE
IZ, a ee CZ
Y : — NE MINwWA“ZZ
Zz < SIX i SS U// —_ SOs =
TS ISS V\\\\Se2
YN NY, Sh / A= } Y\\ \) —s
aN I SNe {= “NS

but the name has been applied rather to irregularly many-times
parted or dissected leaves (such as those of Dicentra), or to
those more than thrice compounded. Of regularly twice or
thrice compound leaves, the commonest are the

Bipinnate or Twice Pinnate, of ordinary

occurrence in the Mimoseous and Cvesalpi- ;
neous, but not in the Papilionaceous, Legu- a
minose. Fig. 208 represents a bipinnate leaf if y

of the Honey Locust (Gleditschia), with the
variation (common with that tree) that some
of the partial petioles, in this figure only the
lowest, bears a single leaflet, while the others 7 ‘
are extended into secondary rhachises fur- }

nished with numer- ue
ous leaflets, mostly cS e pie
j : > pin- <—Sdioapsse /) ;
in the abruptly pin- <Sy ie: 5

iN
te style. On th = 14 = WOH H i \
ae Bee the are = (I Uijz= = i H y
leaves, which are iy! // “is | ; : U
lustered hort <7 =: J
clustered on sho =i lS

spurs, are simply
pinnate. The large
leaves of Gymnocladus are similarly and abruptly bipinnate,

FIG. 208. A bipinnate and multifoliolate leaf of Gleditschia or Honey Locust.
FIG. 299. Bipinnate leaves of Sensitive Plant, Mimosa pudica, with approximate
pinne.

104 MORPHOLOGY OF LEAVES

except at the base, which is simply pinnate or with one or two
pairs of simple leaflets.

Tripinnate or Thrice Pinnate leaves of a regular sort are rare ;
but, with some irregularity, they occur in many species, as in
Aralia, &c. This extent of division, and even much greater, is
common in Ferns.

Digitate-Pinnate is where the primary division of the petiole is
on the palmate or digitate plan; the secondary, on the pinnate.
This seems to be the case in the Sensitive Plant, Mimosa pudica,
Fig. 209. But the leaf is here truly bipinnate with the primary
divisions very crowded at the apex of petiole.

Conjugate-Pinnate is the same arrangement, with the primary
divisions a single pair, at the apex of the petiole, and the leaflets
pinnately arranged on these.

Digitately or Palmately Decompound in a nearly reel way
is not an uncommon case.
Usually, the petiole is succes-
sively three-forked, as in Fig.
210, when the leaf is said to
be biternate (twice ternate),
triternate (thrice ternate), or
quadriternate (four times ter-
nate), &c., according to the
number of times it divides, or
2-3—4-times ternately compound.

an The ultimate divisions in such
cases of threes are commonly of the pinnately trifoliolate type.

201. Pinne is a convenient name for the partial petioles of a
bipinnate leaf, taken together with the leaflets that belong to
them. Thus, the Sensitive Plant, Fig. 209, has four pinne, or
two pairs; the Honey Locust, Fig. 208, a greater number.
When such leaves are still further compounded, the pinn of
higher order, or the ultimate ones, take the diminutive term of
PrxnuL& or Pixnutes. ‘The blades these bear are the Leaflets.

202. The Petiole or Leafstalk is a comparatively unessential
part of the leaf. It is often wanting (then the blade is sess¢le) ;
it may be absent even in compound leaves of the palmate type,
the leaflets rising side by side from the stem. When present, it
is usually either round, or half-cylindrical and channelled on the
upper side. In the Aspen, it is flattened at right angles with
the blade, so that the slightest breath of air puts the leaves in
motion. Sometimes it is much dilated and membranaceous at

FIG. 210. Quadri-ternately compound or ternately decompound leaf of Thalictrum
Cornuti.

THELR STRUCTURE AND FORMS. 105

base, as in many umbelliferous plants; sometimes it forms a
sheath, occasionally it is bordered with appendages, &c. Peti-
oles may assume special functions, to be hereafter considered.
The woody and vascular tissue runs lengthwise through the
petiole, in the form usually of a definite number of parallel
threads, to be ramified in the blade. The ends of these threads
are apparent on the base of the leafstalk when it falls off, and on
the scar left on the stem, as so many round dots (Fig. 81, 85,
91), of a uniform number and arrangement in each species.

203. Partial Petioles are the divisions of the petiole in a
compound leaf. The footstalk of a leaflet takes the diminutive
name of PETIOLULE.

204. Stipules (157) are lateral appendages, one each side of
the base of the petiole, sometimes free
from it and from each other (Fig.
142), sometimes attached by one
edge to its base (Fig. 211), some-
times united with each other into a 4
single body (Fig. 212) in various
ways or degrees. In the latter case,
they usually appear to be within the
base of the leaf or leafstalk; or, as
in the Plane-tree, they may be joined
into one over against the leaf, as
if opposite to it, but their normal
position is supposed to be lateral or
marginal to the petiole. Sometimes
they are foliaceous in appearance and
in function; sometimes they are dry
and colorless or scale-like, reduced to
mere epidermal tissue, and evidently
functionless ; sometimes (as in Mag-
nolia, Fig. 81, Fig-tree, and Beech),
they serve as bud-scales, and fall when
the leaves develop ; sometimes they are
reduced to a mere bristle, or take the a a
form of a spine, as in the Locust (Robinia). Between salient
expansions or wing-like margins of the base of the petiole,
such as those of the Saxifrage tribe, and stipules adnate to the
margins of the petiole, as in most Rosacew, there is no clear
limitation. But presence or absence of stipules generally runs

FIG, 211. Clover-leaf, with adnate stipules. 212. Ochreate stipules (ochrea or ocrea!
of Polygonum orientale, sheathing the stem for some distance, and ending in a spread-
ing border.

106 MORPHOLOGY OF LEAVES.

through a naturai order. Yet what are called stipules in one order
may pass for expansions or appendages of the petiole in another.
In Spergularia, some stipules are connate around the base of the
pair of leaves, including them as well as the stem in the sheath."

205. Stipules, which are normally a pair, may unite into one
body, either adnate to the inner face of the leaf, as in some
species of Potamogeton, or united opposite the leaf, as in Plane-
tree, or united znfer se in a sheath, as in Polygonum. Also when
the leaves are opposite and the stipules thus brought into prox-
imity, the adjacent half stipules of the two leaves may coalesce,
and present the appearance of only two stipules to two leaves,
as in many Rubiacez. A notch or fork at the apex often
indicates the composition.

206. Sheathing stipules, like those of Polygonum (Fig. 212),
are said to be oehreate, or (better) ocreate ; the sheath, thus
likened to a leggin or the leg of a boot, is an OcureEa, as written
by Willdenow, or better Ocrra.

207. The Licute of Grasses (Fig. 150) is seemingly a thin and
scarious extension of the lining to the sheath which answers to
petiole in such leaves: it projects at the junction of the sheath
and blade, there forming a kind of ocrea; and it is generally
regarded as a sort of stipule.

208. Stipels (Stipella) are as it were stipules of leaflets, which
are common in certain tribes of Papilionaceous Leguminosee, e. g.
in the Phaseolex, in Wistaria, Locust. &c.; also in Staphylea.
They are small and slender, and, unlike stipules, they are single
to each leaflet, except to the terminal one, which has a pair. As
leaves furnished with stipules are said to be st/pulate, so leaflets
with stipels are st/pellate.

209. Some unusual modifications of leaves as foliage. In
leaves as illustrated thus far, it is the lamina or blade which is
expanded to do the work of foliage; which is expanded hori-
zontally, so as to present upper and under surfaces, one to the
sky, the other to the ground; which is bilaterally symmetrical
or substantially so, the two lateral halves being nearly if not
quite alike ; and which is affixed to the stem at the basal margin,
or some part of it, with or without a petiole. Various deviations
or apparent deviations from this pattern occur. Some of them are
of comparatively small account and simple explanation, such as

216. Inequilateral Leaves, being unsymmetrical by the much
greater development of one side. This is illustrated in the
whole genus Begonia (as in Fig. 161), consisting of many spe-

! As pointed out by Prof. A. Dickson, in Nature, xviii. 507.

THEIR STRUCTURE AND FORMS. 107

cies, some of which are moderately, and most of them strikingly,
oblique in this way. Elm-leaves, and the like, are more or less
inzequilateral at the base.

211. Connate and Perfoliate Leaves. These are explained by
the union of con-
tiguous leaf-edges.
Peltate leaves, to
which a paragraph
has already been
given (182), come
under the same
head; the seeming
attachment of the
petiole to the lower
face of the blade
being the result of
a congenital union
of the edges of the
sinus. In a sessile ?
leaf, when such a 213 a
union takes place, it surrounds and encloses that portion of the
stem ; which is thus perfoliuate. (Fig. :
213, 214.) It is the stem which is
literally perfoliate, ¢.e. which seem-
ingly passes through the leaf; but it
is customary, though etymologically
absurd, to call this a perfoliate leaf!
Uvularia perfoliata (Fig. 213), in the
later growth of the season, reveals the
explanation of the perfoliation: the
base of the lower leaves conspicuously
surrounds and encloses the stem: that
of the upper is merely cordate and
clasping ; the uppermost simply ses-
sile by a rounded base. Baptisia
perfoliata (Fig. 214) is a more
strongly marked case of perfoliation.
But there are good morphological
reasons for inferring that this seemingly simple leaf consists of
a pair of stipules and a leaflet combined. An occasional mon-
strosity verifies this supposition.

WM
KW \ a

FIG. 213. Leafy branch of Uvularia perfoliata.

FIG. 214. Leafy and flowering branch of Baptisia perfoliata.

FIG. 215. Lonicera flava, a wild Honeysuckle, connate-perfoliate as to the upper
leaves.

108 MORPHOLOGY OF LEAVES.

212. When leaves are opposite, the perfoliation (such as that
of Honeysuckles, Fig. 215) is obviously the result of a congeni-
tal union of the bases of the pair by their contiguous edges.
Leaves connate in this way by narrow bases are not rare nor
remarkable ; but when the two are thus coalescent into one broad
foliaceous body, giving this appearance of perfoliation, the term
connate-perfoliate is used to express it.

213. Vertical Leaves, those with blades of the ordinary kind,
but presenting their edges instead of their faces
to the earth and sky, or when erect with one
edge directed to the stem and the other away
from it, are not uncommon. They prevail in
the Australian Myrtacez, &¢., and occur with
less constancy in the Californian Manzanitas,
and in a great variety of herbs and shrubs. The
anomaly involves no exception to the rule that a
leaf-blade is always expanded in the horizontal
plane, when expanded at all; for, except in equi-
tant leaves, it is the result of a twist of the petiole
or of the blade itself.! In strongly marked
cases, or in most of them, the organization of
the epidermis and superficial parenchyma and
the distribution of the stomata are the same on
both faces.

214. Equitant Leaves are vertical on a different L\'t
plan. They are conduplicate, ¢.e. are folded 6 * ¥

&>

UMN

E> yy

217 26
together lengthwise on their middle, the upper surface thus con-
cealed within, the outer alone presented to the air and light.

atta as
i

1 Silphium laciniatum, the so-called Compass Plant, and (hardly less so)
S. terebinthinaceum, are good instances of the kind, most of the leaves
making a half-twist, the radical ones by their long petioles. In the former
species, the pinnately parted blade occasionally makes a farther twist, so as
to bring the upper part into a plane at right angles to the lower. The
blades place themselves in various directions as respects the cardinal points ;
but on the prairies the greater number affect a north and south direction of
their edges,—a peculiarity first pointed out, in the year 1842, by General
B. Alvord, U.S. A.

FIG. 216. Equitant erect leaves of Iris, with the rootstock.
FIG. 217. A section across these leaves at the base, showing the equitant character.

THEIR STRUCTURE AND FORMS. 109

Being two-ranked and closely crowded, the outer ones at their
base fold over or bestride the inner (as shown in the sectional
diagram, Fig. 217), whence the name of eguitant. Above, the
contiguous halves of the inner face congenitally cohere, and so
produce the sword-shaped or linear vertical blade which is
characteristic of Iris (Fig. 216) and the Iris family. In most
there is a farther complication, of an excep-
tional kind, viz. the development backwards
of a portion of blade from the midrib, often
forming most of the upper part of such leaves,
which therefore may really be said to develop
in the vertical plane.

215. Leaves with no distinction of Parts, 7. e.
of blade and petiole. ‘This is the case in Iris
(Fig. 216), Daffodil, the Onion, and perhaps
of most parallel-veined leaves of Endogens.
Those expanded in the horizontal plane may
however be regarded as sessile blades: those
which are not expanded, but filiform, or needle-
shaped (acicular), or awl-shaped (subulate), may
be regarded either as homologous with petioles,
or as unexpanded blades, which amounts nearly a4
to the same thing where there is no trace of a petiole at base.
Under this head may be ranked the leaves of Pines (Fig. 248) ;
also both the subulate and the
scale-shaped and adnate leaves
of Arbor Vite, Red Cedar (Juni-
perus Virginiana), and other trees G
of the Cypress tribe. (Fig. 218.) ~~

216. Stipules serving for Blade.
Lathyrus Aphaca is a good in-
stance of this (Fig. 219); the
petiole becoming a tendril, the
leaflets which its relatives bear
being wholly wanting, the ample
foliaceous stipules assume the
appearance of leaves. In some on
other species of Lathyrus, and in the Pea, equally large stipules
share with the pair or pairs of leaflets in the functions of foliage.
On morphological evidence, we judge that the singular leaves of

FIG. 218. A twig of Arbor Vite, with both awl-shaped and scale-shaped leaves.

FIG. 219. Lathyrus Aphaca: portion of stem, bearing a single leaf, which consists
of a pair of foliaceous stipules, and a petiole in the form of a tendril; in its axil a
flower-stalk.

110 MORPHOLOGY OF LEAVES.

Baptisia perfoliata, shown in Fig. 214, are not simple blades,
but each a pair of stipules, with or without a terminal leaflet, all
completely confluent into one body. The related species of the
genus have trifoliolate leaves and foliaceous stipules ; hence these
simple leaves without stipules are best explained in this way.

217. Phyllodia, or Petioles serving for Blade. Sometimes the
petiole develops foliaceous margins, or wings, as in the Bitter
Orange and in Rhus copallina. These are efficient as foliage in
proportion to their size. These are not to be confounded with the
case in which a petiole specially develops as a blade-like organ, -
which usurps the office of foliage. A petiole-blade of this kind
is named a Puyttopium. Occurring only in Exogens, phylodia
are generally distinguished from true blades by the parallel
venation, and always by their normally vertical dilatation ; 7. e.
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 300 species. The
true lamina of these is bipinnate. It appears on seedlings, and
occasionally on later growths. Several South American species of
Oxalis produce phyllodia. So likewise do our tubular or trumpet-
leaved species of Sarracenia in that portion of the foliage which
develops the pitcher imperfectly, or not at all. Indeed, all
Sarracenia-leaves are phyllodia with the back in most of them
hollowed out into a tube or pitcher; and the terminal hood
answers to the blade.

§ 3. Leaves servine SpEcIAL OFFICEs.

218. Leaves may serve at the same time both their ordinary
and some special use, or even more than one special use. For
example, in Nepenthes (Fig. 222) there is ‘a well-developed
blade, usually sessile, which serves for foliage, a prolongation of
its tip into a tendril, which serves for climbing, then an extraor-
dinary dilatation and hollowing of the apex of this into a pitcher
for a very special use, and a peculiar development of the apex
of this into a lid, closing the orifice during growth. Among the
special purposes which leaves subserve, and the study of which
connects singularities of morphology with teleology, the most
remarkable is that of

219. Leaves specialized for the Utilization of Animal Matter.
This occurs in leaves which also assimilate, or do the ordinary
work of vegetation ; and the special function is usually taken up
by some particular portion of the organ. The details of this

LEAVES SERVING SPECIAL OFFICES.

eet

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

all historical references are relegated. Only the morphology of

such leaves is here under consideration.

220. As Ascidia or Pitchers, vessels for maceration, &c. These

occur in several widely different
families of plants. The commonest
are those of the Sarracenias, natives
of Atlantic North America. They
are evidently phyllodia (217), the
cavity being a hollowed dorsal por-
tion: the wing-like or foliaceous por-
tion, always conspicuous and forming
the ventral border, makes the whole
organ or most of it in the earlier
leaves of the tubular species. The
pitchers of S. purpurea (Fig. 221,
225), the only species which extends
north of Virginia, are open cups,
half filled with water, much of which
may be rain, in which abundance of
insects are usually nndergoing macer-

ation. InS. variolaris (Fig. 226), the hooded summit, answer-
ing to the blade of the leaf, arches over the mouth in such wise

FIG. 220. Pitchers of Heliamphora; 221. of Sarracenia purpuren;

222. of Nepenthes.

223. A phyllodium of a New Holland Acacia. 224. The same, bearing a reduced com-

pound blade.

FIG. 225. Pitcher-leaves of Sarracenia purpurea; one of them with the upper part

cut away.

ya Wy MORPHOLOGY OF LEAVES.

as to mostly exclude the rain; in 8. psittacina (Fig. 227) the
inflexed and inflated hood completely excludes it. The water
which these contain is undoubtedly a secretion.
All entrap flies, ants, and various insects, which
in most species are lured into the pitcher by a
sweetish secretion around or at some part of the
orifice.t Few that have entered ever escape ;
most are decomposed at the bottom of the cavity.
In Darlingtonia Californica (Fig. 228), the Cali-
fornian representative of Sarracenia, the inflated
hood guards against all access of rain, while the
orifice is freely open to flying insects from be-
neath; and a singular two-forked appendage, like
to a fish-tail (probably the homologue of the
blade), overhangs the front. The inner face of
this appendage is besmeared with the sweet and
viscid secretion which allures insects to the open-
ing. In this and in Sarracenia variolaris, the
sweet secretion in the early season is continued
upon the edge of the wing, forming a saccharine
trail which leads from near the ground up to the
orifice of the pitcher.? Fig. 220 represents pitchers
of Heliamphora, a little-known South American
representative of Sarracenia. Its wing is narrow
and inconspicuous, the mouth widely open and
directed upward, and the hood reduced to a
minute and upright, probably functionless ap-
226 pendage. In Cephalotus —an anomalous plant
of Australia, of uncertain affi-
nity — the leaves for foliage
are dilated phyllodia ; among
them are others completely
transformed into stalked and
short pitchers, with thickened
rim and a_ well-fitting lid,
hinged by one edge, Fig. 229. «
The particular morphology of the parts is not well made out.

1 This sweet secretion, which at times is very obvious in the southern
species, has also been detected by Mr. Edward Burgess in S. purpurea; but
it is rarely seen, and probably plays no important part in the capture and
drowning of the multitude of insects which these pitchers are apt to contain.

2 This trail was discovered by Dr. J. H. Mellichamp, of South Carolina.
See Proce. Am. Association for Advancement of Science, xxiii. 113 (1871).

FIG. 226. Pitcher of Sarracenia variolaris. 227. Same of S. psittacina.

LEAVES SERVING SPECIAL OFFICES. 143

221. The pitcher-bearing leaf in Nepenthes has been referred
to (218, Fig. 222): of this there are various species, all of them
somewhat woody climbing
plants of tropical Asiatic
and African islands of
the southern hemisphere,
some of them familiar in
conservatory cultivation.
Here the tendril may be
regarded as a prolonged
extension of the midrib
of the blade, and the
pitcher, with its hinged
lid, as a peculiar development from. as apex. The
water contained in the pitcher is a secretion, much
of which appears before the lid opens ; and a sweetish
excretion at the orifice lures insects. The presence
of these in the pitcher increases the watery secre-
tion in which the animals are drowned; and this
secretion is ascertained to have a certain digestive
power.’

222. The aquatic sacs of Utricularia or Bladder-
wort are diminutive ascidia, always under water, 228
and with lid opening inward, like a valve, preventing the exit
of minute animals entrapped therein.*? Morphologically, they
are doubtless leaves or parts of leaves.

223. As Sensitive Fly-traps. The leaves of all species of
Drosera or Sundew are beset with stout bristles tipped with a
gland, which secretes and when in good condition is covered
by a drop of a transparent and very glairy liquid, sufficiently
tenacious to hold fast a fly or other small insect. Adjacent
bristles, even if not touched, in a short time bend towards those
upon which the insect rests, and thus bring their glands also
into contact with it. In Drosera filiformis, the leaves are fili-
form, with no distinction of petiole and blade. In D. rotundifolia
and other common round-leaved species, there is a clear distinc-

1 This was first made out by J. D. Hooker, and announced in his address,
as President of the British Association for the Advancement of Science, at
Edinburgh, 1871.

2 Darwin, Insectivorous Plants, 395. Cohn, Beitrige zur Biologie der
Pflanzen, 1875. Mrs. Treat, in The Tribune, New York, September, 1874,
and Gard. Chron. 1875, 303.

FIG. 228. Pitcher of Darlingtonia Californica. 229. Pitcher of Cephalotus follicu-
laris, with lid open.

114 MORPHOLOGY OF LEAVES.

tion of petiole and blade, and the stalked glands thickly beset
the whole upper surface of the latter. A small insect alighting
thereon is helpless, and is soon touched by all the glands within
reaching distance; also the blade itself commonly ineurves,
taking part in the general movement. It has recently been
demonstrated that the captured insect is fed upon, and that the
plant thereby receives nourishment. Here leaves which do the
normal assimilative work of vegetation, but somewhat feebly
(having a comparatively small amount of chlorophyll), have

A
we

also the power and the habit of obtaining ready-organized food
by capture, and are benefited by it.

224. Species of Drosera inhabit most parts of the world, and
the genus is numerous in species. A near relative, Dionza, is
of a single species, D. muscipula (Venus’s Fly-trap), inhabiting
only a limited district in the sandy eastern border of North
Carolina. It is more strikingly sensitive and equally carnivo-
rous, but in a different way. It is destitute of stalked and viscid
glands. The apparatus for capture and digestion is the two-
valved body at the top of each leaf. (Fig. 230, 231.) If this

FIG. 230. A plant of Dionwa muscipula, reduced in size. 231 Three of the leaves,
of almost the natural size; one of them open, the others closed. Probably a fly is never
caught by the teeth, in the manner here represented.

LEAVES SERVING SPECIAL OFFICES. de 5)

be taken for the leaf-blade, the part below would be a broadly-
winged foliaceous petiole. If the latter be the true blade, the
apparatus in question must be reckoned as a peculiar terminal
appendage. Both are moderately green, and act as foliage.
The specially endowed terminal portion acts also in a decidedly
animal-like manner. When either of the three or four slender
bristles of the ‘upper surface are touched, the trap suddenly
closes, by a movement ordinarily quick enough to enclose and
retain a fly or other small insect. The intercrossing of the stout
marginal bristles detains the captive, unless it happens to be
small enough to escape by the intervening little openings.
Otherwise, the sides soon flatten and are brought firmly into
contact, and a glairy secretion is poured out from numerous
immersed glands: this, with the extracted juices of the macerated
insect, is after some time reabsorbed; the trap, if in a healthy
condition, now re-opens and is ready for another capture. For
references to the now copious literature of this whole subject, and
for its physiological treatment, the succeeding volume should be
consulted.

225. Leaves for Storage. Nutritive matter is stored in leaves
in many cases, and not rarely in leaves which at the A pT
same time are subserving the purpose of foliage. / //// f\\
This occurs in all fleshy leaves, to a greater or less
extent, according to the degree of thickening or
accumulation. The leaves of the Century Plant
or Agave, for instance, are green and foliaceously
efficient at the surface, while the whole interior is
a store-house_of farinaceous and other nutritious
matter, as much so as is a potato. The leaves
_of various species of Aloe, Mesembryanthemum,
Sedum, and other ‘‘ succulent” plants (in which
a large part of the accumulation is water) are not
rarely so obese as to lose or much disguise the
foliaceous appearance. Sometimes one portion of
a leaf is of normal texture and use, while another
is used as a reservoir for the nourishment which the
foliaceous part has produced. Fig. 252, a leaf from
the bulb of White Lily, the base of which forms
one of the bulb-scales, is an instance of the kind.! The most

1 In Dicentra Cucullaria and (more strikingly from the sparseness of the
grains) in D. Canadensis, the matter elaborated in the much dissected blade is
conveyed to the very base of the long petiole, and there deposited in a con-

FIG. 232. A radical leaf of the White Lily, with its base thickened into a bulb-scale,
which is cut across to show its thickness.

116 MORPHOLOGY OF LEAVES.

decisive instance of leaves used for storage of food is in that
material provision for the nourishment of the embryo in germi-
nation, in which the first leaves, the cotyledons, are turned to this
account. (21-37, &c.) After or
while discharging this special duty,
the cotyledons may fulfil their gen-
eral office, by serving as foliage (as
in Maples, Fig. 8, and Pumpkins,
Fig. 47) ; or, through various inter-
mediate conditions, they may be
wholly devoted to storage, as in
the Pea, Oak, Horsechestnut, &e.
(Fig. 37-43.)

226. Leaves as Bulb-seales, how-
eyer, are for the most part wholly
applied to this use, being leaves
reduced to short scales or to
concentric coats, and thickened
throughout by nutritive deposit.
The accumulation of such leaves
forms the mass of the bulb, as of
the Lily, Fig. 118, Onion, Fig.
113, &e., also of bulblets. (120.)

227. Leaves as Bud-seales, being
for protection of nascent parts, have
been explained under buds. (70.)
The evidence of foliar nature af-
forded by transition is well exhib-
ited by the Sweet Buckeye, although
the whole series of gradations, from
bud-scales to compound leaves,
is seldom seen united in one bud,
as in Fig. 233. In this case, the
bud-seales are homologous with
petioles. In Magnolia, they consist
of stipules (Fig. 81, 82): in the
Lilac, they are homologous with leaf-blades. The two pairs of
bud-seales which subtend and protect through winter the nascent
head of flowers of Cornus florida are morphologically the apex of

centrated condition, in the form of a solid grain, which remains for next
year’s use, the whole leaf except this thickened base dying away at the close
of the short season’s growth.

FIG 233. Leaves of a developing bud of the Low Sweet Buckeye (#sculus parvi-
flora), showing a nearly complete set of gradations from a scale to a compound leaf ot
five leaflets.

LEAVES SERVING SPECIAL OFFICES. 417

blades. When the blossoms develop in spring, these scales grow
from beneath, greatly expand, and become obovate or obcordate
petaloid leaves, the brown terminal notch
of which is the bud-scale, which was un-
able to take part in the vernab growth.

227%. Leaves as Spines. All gradations
may be found between spiny-toothed leaves
(as in Holly), in which teeth are pointed and
indurated, and leaves which are completely
contracted into a simple or multiple spine.
Indeed, such a transition is seen in the Bar-
berry, Fig. 254. The foliar nature of such
spines is manifest from their position, sub-
tending a bud from which the foliage of the
season proceeds, and themselves not sub-
tended by any organ. In some Astragali,
the petiole of a pinnate leaf indurates into
a slender spine and persists, the leaflets
early falling. The spine in Fouquiera is a
portion of the lower side of the petiole or
midrib, indurated and persistent, the rest
of the leaf separating by splitting when it
has served its office.!

228. Leaves adapted to Climbing. Some plants climb by the
action of the stem or of certain branches specially ue to
this purpose (99): others gain the needful
support by means of their leaves (101) ; some-
times by an incurvation of the tips, either of
a simple blade as in Gloriosa, or small partial
blades, as in Adlumia, and often in Clematis,
thereby grappling the support ; sometimes by
the petiole making a turn or two around a
support (as in Maurandia, climbing Antirrhi-
nums, Rhodochiton, and Solanum jasminoides, §
Fig. 235) ; sometimes by the transformation f4
of one or more leaflets of a compound leaf 7 ¥
into tendrils, as in the Pea and Vetch (Fig.
204) ; sometimes by the suppression of all the
leaflets and the conversion of the whole petiole into a tendril, as
in Lathyrus Aphaca (Fig. 219) ; and perhaps by the conversion

234

1 Described in Plante Wrightiane, ii. 65.
FIG. 234. A vernal shoot of common Barberry, showing a lower leaf in the normal
state; the next partially, those still higher completely, transformed into spines.
FIG. 235. Solanum jasminoides, climbing by coiling and at length indurating petioles.

118 MORPHOLOGY OF LEAVES.

of a pair of stipules into tendrils in Smilax. At least the ten-
drils here occupy the position of adnate stipules. The tendrils
of Cucurbitaceze are peculiar and ambiguous, on account of their
lateral and extra-axillary position and the manner in which the
compound ones develop their branches. But they are doubtless
partly if not wholly foliar.’

229. Petaloid Leaves, Bracts. Certain leaves, situated near to
flowers, and developing little or no chlorophyll in their paren-
chyma, exchange the ordinary green hue and herbaceous texture
for the brighter colors and more delicate structure which are
commonly seen in and thought to characterize flower-leaves.
Such are said to be colored, meaning, as applied to foliage, of
some other color than green. As petals are the type of such
colored parts, they are said to be petalord, i. e. petal-like. They
are like petals, moreover, in one of the purposes which these sub-
serve. (299.) Examples of these petaloid leaves are seein in the
shrubby Mexican Euphorbia called Poinsettia, in Salvia splen-
dens, most species of Castilleia or Painted Cup, also in the
white hood of Calla and Richardia /thiopica (called Calla Lily),
and in the four white leaves which subtend the flower-head of
Cornus florida, and of the low herbaceous Cornel, C. Canaden-
sis. (Fig. 294.) Such leaves, being in proximity to flowers, and
all others which are within a flower-cluster or are borne by
flower-stalks, receive the special name of Bracrs. More usually
bracts are not petaloid, but different in size or shape from ordi-
nary leaves, either by abrupt change or gradual transition. Not
uncommonly they are reduced to scales or mere rudiments or
vestiges of leaves, of no functional importance.

230. Flower-Leaves. The morphology of leaves extends not
only to ‘** the leaves of the blossom,” more or less accounted as
such in common parlance, but also to its peculiar and essential
organs, the relation of which to leaves is more recondite. Their
morphology needs to be treated separately, and to be preceded
by a study of the arrangement of leaves and of blossoms.

1 The most satisfactory interpretation may be that of Braun and Wydler,
adopted by Eichler (Bliithendiagramme, i. 304): that the flower of Cucur-
bita and its peduncle represent the axillary branch, the tendril by its side
answers to one of the bractlets (that of the other side being suppressed),
and the supernumerary branch springs from the axil of the tendril. This
makes of the tendril a simple leaf, of which the branches are the ribs. But
the tendril-divisions are evidently developed in spiral order, and in vigorous
growths occupy different heights on the tendril-axis. This favors Naudin’s
view, that the main tendril is cauline, and its divisions leaves.

PHYLLOTAXY, OR LEAF—ARRANGEMENT. 119

CHAPTER IV.

PHYLLOTAXY, OR LEAF-ARRANGEMENT.

Section I. THe DIstrisuTION oF LEAVES ON THE STEM.

231. Puytiotaxy (or Phyllotaxis) is the study of the distri-
bution of leaves upon the stem and of the laws which govern it.
The general conclusion reached is, that leaves
are distributed in a manner to economize space
and have a good exposure to light, &c., and
that this economy on the whole results from
the formation of leaves in the bud over the
widest intervals between the
leaves next below.! Leaves
are arranged in a consider-
able variety of ways, which
all fall under two modes, the (==>
Verticillate and the Alter-\G
nate (13), but which may
also be termed the Cyclical g
and the Spiral. f

232. Alternate leaves are
those which stand singly,
one after another; that is,
with one leaf to each node i
or borne 236 237
on one height of stem. Verticillate leaves are
those with two or more at the same height of
stem, circularly encompassing it, 7. e. forming:
a Verticil or Whorl. Verticillate and whorled
are synonymous terms to denote this arrange-
ment. These two Kinds of leaf-arrangement
> are commonly ranked as three, viz. alternate,
opposite, and whorled. But the opposite is
only the simplest case of the whorled, being

1 For the most comprehensive discussion of phyllotaxy in connection with
development, and in view of these relations, see Hofmeister, Allgemeine
Morphologie, § 11, and Chauncey Wright, Mem. Amer. Academy, ix. 389.

FIG. 236, Alternate, 237, Opposite, 238, Verticillate or whorled leaves.

120 PHYLLOTAXY, OR LEAF-ARRANGEMENT.

that in which the members are reduced to two. ‘This case is
so much commoner than whorls of three and of higher numbers
that it took from the first its special name of opposite, so that in
descriptions the phrase ‘‘ leaves verticillate ” implies more than
two leaves in the whorl. But it should be kept in mind that
“leaves opposite ” is the same as ‘* leaves in whorls of two.”

233. The greater number of phanogamous plants (all but the
monocotyledonous class) begin with verticillate leaves, mostly
of the simplest kind (7. e. cotyledons opposite) : some continue
verticillate throughout; some change in the first leaves of the
plumule or after the first pair into alternate, and again into
verticillate in or toward the blossom, in the interior of which the
alternate arrangement may be again resumed. As Nature passes
readily from the one mode to the other on the same axis, we
may expect that the two may be comprised under some common
expression. But they have not yet been combined, except by
gratuitous or somewhat forced hypotheses; so that for the
present they should be treated in morphology as primarily dis-
tinct arrangements.!

254. Verticillate or Cyelical Arrangement. Here the leaves
occupy a succession of circles, or form whorls around the stem,
two, three, four, five, &c., in each whorl. According to the
number, the leaves are opposite, ternate, quaternate, quinate, and
so on. The characteristic of the individual whorl is that the
members stand as far apart from each other as their number
renders possible, 7. e. they divide the circle equally. Thus, when
only two, or opposite, their midribs or axes of insertion have an
angular divergence (as it is termed) of 180°; when three, of
120°; when four, 90°; when five, 72°.

255. The characteristic of the whorls mn relation to each other
is, that the members of successive whorls stand over or under
the intervals of the adjacent ones. In other words, successive
whorls alternate or decussate. ‘This economizes space and light,
or gives the best distribution which the cyclical system is capa-
ble of. And it is in accordance with the general conclusion of
Hofmeister’s investigation of the origin of phyllotaxic arrange-
ments in the nascent bud, viz. that new members originate just
over the widest intervals between their predecessors next below.
Thus, in opposite leaves or whorls of two (Fig. 237), the suc-
cessive pairs deeussate or cross at right angles, and so four

1 It is readily seen that whorls may be produced by the non-development
of the internodes between the leaves of a series of two, three, five, or more
in alternate order. The difficulty is that the members of the next whorl do
not follow the order that they should upon this supposition.

DISTRIBUTION OF LEAVES ON THE STEM. 121

straight equidistane vertical ranks are produced. In ternate or
trimerous whorls there are six vertical ranks; in quaternate or
tetramerous whorls, eight vertical ranks, and so on.!

236. The cases in which successive ‘S)
pairs of leaves do not decussate at right ———

angles, or the members of whorls are not Fe
exactly superposed to intervals, but as it

were wind spirally (as in Dipsacus, many f
Caryophyllacez, &c.), may some of them ee y
be explained by torsion of the stem, So

such as is very manifest in numerous in- %

SS

stances ; and others may be resolved into 239
instances of alternate leaves simulating or passing into whorls
by the non-development of internodes.?

237. Alternate or Spiral Arrangement. Here the leaves are
distributed singly at different heights of the stem, and at equal
intervals as respects angular divergence. (Fig. 256.) This angu-
lar divergence (/.e. the angular distance of any two successive
leaves) differs in the various kinds of this system of phyllotaxy,
but is always large enough to place the leaves which immediately

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

2 In Lilium Canadense, superbum, &¢., with whorls of variable number
of leaves and vague relation to each other (when of the same number some-
times the members superposed), and above and below passing into the alter-
nate arrangement normal to the family, these whorls are evidently formed
of alternate leaves brought together by non-development of internodes.

Here may also be mentioned the not uncommon anomaly in Fir-cones,
notably those of Norway Spruce, the normal phyllotaxy of which is simply
spiral, but in occasional instances the cone is composed of pairs of opposite
scales, spirally arranged, 7. e. the pairs not decussating at right angles, thus
forming double spirals. In the abnormal spruce-cones, the fractions usually
observed are 7 or 7g, or, as expressed by Braun, (4)35 and (4)3°s.

Braun’s mode of notation for the ordinary succession (?. e. the decussation )
of opposite leaves is (4)4, the 4 meaning that the two leaves of the pair are
half the cireumference of the circle apart, the denoting that each leaf of
the succeeding pair diverges one fourth of the circumference from the pre-
ceding. Braun finds cases in which pairs (and equally whorls) are super-
posed (e.g. certain species of Mesembryanthemum and Euphorbia). these
are expressed in this notation by the formula (4)4, that is, the corresponding
leaves of the succeeding pair diverge 180° from their predecessors. He
recognizes also some cases of intermediate divergence; such as (4)? in the
upper leaves of Mercurialis perennis, (}),°; on certain stems of Linaria vul-
garis, (4)3°; exceptionally in the leaves of Epilobium angustifolium and the
scales of Norway Spruce, (4)3 exceptionally in the scales of Norway
Spruce. See Ordnung des Schuppen an der Tannenzapfen, 376, &c.

FIG. 239. Ground-plan diagram of six trimerous whorls, showing their alternation.

122 PHYLLOTAXY, OR LEAF—ARRANGEMENT.

follow each other in the ascending order upon different sides of
the axis: it also secures an advantageous spacing of the leaves
over the whole length of the axis. Their vertical distance from
each other of course depends on the length which the internodes
attain, which is a matter of growth and is very variable; but
their angular distance is fixed in the kind or numerical plan of
the particular phyllotaxy, and is uniform throughout.

238. The leaves are said to be alternate, because they come
one after another, now on this side, then on that, as they ascend
the stem. The arrangement is said to be spzral, because if a line
be drawn or a thread extended from the base or insertion of one
leaf to that of the next higher, and so on, taking in all the leaves,
it forms a helix, more or less loose or close according to the
development of the internodes. (See Fig. 242.) This imagined
spiral line ascends continuously, without a break; and on it the
leaves are equably laid down.!

239. Almost all the ordinary instances of spiral phyllotaxy
belong to one series, having very simple arithmetical relations.
So that this may be taken as the type, and the few others re-
garded as exceptions or sometimes as modifications of it. The
kinds are simply designated by the number of vertical ranks of
leaves: they are technically named by prefixing the proper
Greek numeral to the word meaning row orrank. The arrange-
ment called

Distichous, or Two-ranked, is the simplest and among the com-
monest, occurring, as it does, in all Grasses and many other
monocotyledonous plants, in Lindens, Elms, and many dico-
tyledonous genera. Here the leaves are disposed alternately on
exactly opposite sides of the stem (as in Fig. 1); the seeond
leaf being the farthest possible from the first, as is the third from
the second ; the third therefore over the first, and the fourth over
the second, and so on, thus forming two vertical ranks. The
angular divergence is here half the circumference, or 180°; and
the phyllotaxy may be represented by the fraction 4, which desig-
nates the angular divergence, while its denominator expresses
the number of vertical ranks formed.

Tristichous, or Three-ranked, is the next in the series, and is

1 But when we reach a leaf which stands directly over a lower and older
one, we say that one set or spire is completed, and that this leaf is the first
of a succeeding set or spire. From analogy of such an open spire to the
closed cycle of a whorl of leaves, it is not unusual to designate the former
likewise as a cycle. Yet it is better (with Eichler) to restrict that term, and
the adjective cyclical, to verticillate phyllotaxy, or to whorls, to which it
properly and etymologically belongs.

—

DISTRIBUTION OF LEAVES ON THE STEM. 123

less common, though not rare in 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 4 designates this
arrangement. ‘Fhe angular
divergence, or distance of
the axis of the first leaf
from the second, and so on,
is one third of the circum-
ference (or 120°): conse-
quently the fourth leaf comes
over the first, the fifth over
the second, the sixth over
the third, and so on; that
is, the leaves fall into three
vertical ranks. The spiral
character here begins to be
manifest, or becomes so by
drawing a line on either fig-
ure from the axis or midrib
of the first leaf to that of
the second, and so on to the
sixth, forming a helix of
two turns.!

Pentastichous, or Five-ranked, sometimes termed the guincuncial
arrangement. This is the most common in alternate-leaved
dicotyledonous plants. It is shown in Fig. 236 (on a branch
of Apple-tree), and by diagrams, displaying the spiral character,
in Fig. 242, 243. The angular distance from the first to the
second leaf (passing the shorter way) is 2 of the circumference,
or 144°. But the spiral line makes two turns round the stem,
on which six leaves are laid down, with angular divergence of 2,

1 The line is supposed to follow the nearest way, and the divergence is
counted as 4, 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 2.

FIG. 240. Piece ofa stalk, with the sheathing bases of the leaves, of a Sedge-Grass
(Carex cruscorvi), showing the three-ranked arrangement. 241. Diagram of the cross-
section of the same. The leaves are numbered in succession.

FIG. 242. Diagram of position of six leaves in the five-ranked arrangement: a spiral
line is drawn ascending the stem and passing through the successive scars which mark
the position of the leaves from 1 to 6. It is made a dotted line where it passes on the
opposite side of the stem, and the scars 2 and 5, which fall on that side, are made
fainter. 243. A plane horizontal projection of the same; the dotted line passing from
the edge of the first leaf to the second, and so on to the fifth leaf, which completes the
turn: as the sixth would come directly before, or within, the first.

124 PHYLLOTAXY, OR LEAF-ARRANGEMENT.

and the sixth is the first to come over any one below; the
seventh comes over the second, the eighth over the third, &e.
The leaves are thus brought into five vertical ranks; but these
7 five leaves are laid down on two turns of the
9 helix (the sixth beginning the second revolu-
> | tion); the angular divergence of the leaves in
; * order is 2, or 144°; the angular distance of
1 © the vertical ranks, 72°. This is a very advan-
tageous distribution for ordinary foliage on
erect or ascending branches. Its formula is 2, expressing the
angular divergence, the denominator also indicating the number
of vertical ranks, the numer-
ator indicating the number
of revolutions made in add-
ing one leaf to each rank.
Vig. 244 illustrates this ar-
rangement on a cone of
American Larch, the scales
of which are homologous
with leaves, the numbers
in sight are affixed, and those of the whole
cone displayed on a plane at the side.
Octostichous, or Hight-ranked, a less common
arrangement, occurs in the Holly, Aconite,
the radical leaves of Plantago. It has the
angular divergence of 135°, or 2 of the cir-
cumference, and the leaves in eight ranks,
the ninth over the first and at the completion
of the third revolution: it is therefore repre-
sented by the fraction 3.
240. The obvious relations of the fractions
3, 1, 2, 2, representing the primary forms of
spiral phyllotaxy, are that the sum of any two
numerators is the numerator of the next suc-
ceeding fraction, and the same is true of the
denominators ; also the numerator is the same
as the denominator of the next but one pre-
ceeding fraction. Following these indications,
the series may be extended to 75, #, 43, 34, &c. Now these

v0

10

=

FIG. 244. A cone of the small-fruited American Larch (Larix Americana), with
the scales numbered, exhibiting the five-ranked arrangement.

FIG. 245. An offset of the Houseleek, exhibiting the 5-13 arrangement; the leaves
in sight numbered, the 14th over the first, the 19th over the 6th, &e.

FIG. 246. Cone of White Pine (Pinus Strobus) with scales numbered from bottom,
and some secondary spirals marked.

DISTRIBUTION OF LEAVES ON THE STEM. #25

eases actually occur, and ordinarily only these.’ The 4% and
#, are not uncommon in foliage. The rosettes of the House-
leek exhibit the 5, or thirteen-ranked arrangement, as also does
the cone of Pinus Strobus, the 14th leaf falling over the first.
(Fig. 246.) The 3 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 3}
and 24 phyllotaxy may be readily made out. This actual series,
4, 4, 2, 2, &c., answers to and may be expressed by the con-

tinued fraction, 44 1

iz ;
Se ter &e.?

1 When other instances are detected, they are found to belong to other
series, following the same law, such as the rare one of }, 4, 3, ;%;.

2 “The ultimate values of these continued fractions extended infinitely
are complements of each other, as their successive approximations are, and are
in effect the same fraction, namely, the irrational or incommeasurate inter-
val which is supposed to be the perfect form of the spiral arrangement.
This does, in fact, possess in a higher degree than any rational fraction the
property common to those which have been observed in nature; though
practically, or so far as observation can go, this higher degree is a mere
refinement of theory. For, as we shall find, the typical irrational inter-
val differs from that of the fraction ? by almost exactly 7755, a quantity
much less than can be observed in the actual angles of leaf-arrangements.”
“On this peculiar arithmetical property . . . . depends the geometrical one,
of the spiral arrangement, 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, according to such an arrangement, no leaf would ever
fall exactly over any other; and, as I have said, we have no evidence, and
could have none, that this arrangement actually exists innature. 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
2 and 3° 3, &e., which are all indistinguishable as measured values in the
plant, do actually realize this property with all needful accuracy. Thus,
§ = 0.375, ~; = 0.385, and 34, = 0.381, and differ from & [the ultimate value
to which: the fractions of this series approximate, or what is supposed to be
the type-form of them] by —0.007, +0.003, and —0.001 respectively ; or they
all differ by inappreciable values from the quantity which might therefore be
made to stand for all of them. But, in putting & for all the values of the
series after the first three, it should be with the understanding that it is not
so employed in its capacity as the grand type, or source of the distributive
character which they have, —in its capacity as an irrational fraction, — but
simply as being indistinguishable practically from those rational ones.” —
Chaucey Wright, in Mem. Amer. Acad. ix. 387-390.

126 PHYLLOTAXY OR LEAF-ARRANGEMENT.

241. The successive grades of angular divergence of alternate
leaves, as expressed in degrees, are
== 180° 144° ps = 138° 27! 41.54"
= 120° 135° #, = 137° 8! 34.29"
and so on; and beyond, if not in the latter cases, the differences
become quite too small for determination by inspection. They
all fall within the } and } as to amount of divergence: and they
form a series converging to a deduced typical angle of 137° 30!
28”, which, being irrational to the circumference, would place
no leaf exactly over any preceding one, but alternately and
more and more slightly on one and the other side of the vertical,
and so on, in an endless spiral. That is, according to Bravais,
the ranks in the higher grades tend to become curvisertal, or
actually become so; while in the lower grades they are obviously
rectiserial. Unless. indeed, there is some torsion of the axis,
by which the vertical ranks are rendered oblique, as is often the
case in cones of the Norway Spruce. But, apart from this, the
difference between rectiserial in a high order and curviserial
soon becomes inappreciable. Any and all of the higher grades,
and practically one as low as the 3, secures the utility of the
theoretical angle, viz., that ‘* by which the leaves would be dis-
tributed most thoroughly and rapidly around the stem, exposed
most completely to light and air, and provided with the greatest
freedom for symmetrical expansion, together with a compact
arrangement in the bud.” Even in the simpler grades of com-
monest occurrence, each leaf (according to Wright) is so placed
over the space between older leaves nearest in direction to it as
always to fall near the middle of the space, until the circuit is
completed, when the new leaf is placed over an old one.!

242. It is to be noted that the distichous or 34 variety gives
the maximum divergence, viz. 180°, and that the tristichous or
4 gives the least, or 120°; that of the pentastichous or 2 is nearly
the mean between the first two; that of the 2, nearly the mean
between the two preceding, &c. The disadvantage of the two-
ranked arrangement is that the leaves are soon superposed and
so overshadow each other. This is commonly obviated by the
length of the internodes, which is apt to be much greater in this
than in the more complex arrangements, therefore placing them
vertically farther apart; or else, as in Elms, Beeches, and the

i ole

Wl
im

1 This corresponds with Hofmeister’s general rule, that “new lateral
members have their origin above the widest gaps between the insertions of
the nearest older members.” Yet the fact that the character of the leaf-
arrangement is laid down at the beginning in the bud does not go far in the
way of the mechanical explanation which he invokcs.

DISTRIBUTION OF LEAVES ON THE STEM. 127

like, the branchlets take a horizontal position and the peti-
oles a quarter twist, which gives full exposure of the upper
face of all the leaves to the light. The } and 2, with dimin-
ished divergence, increase the number of ranks; the 2 and all
beyond, with mean divergence of successive leaves, effect a more
thorough. distribution, but with less and less angular distance
between the vertical ranks.

242°. The helix or primitive spiral upon which the leaves
successively originate ascends, sometimes from left to right,
sometimes from right to left,’ commonly without change on the
same axis, and prevailingly uniform in the same species; but
occasionally both directions occur in the same individual. The
earliest leaves of a stem or branch, or the last, are often on a
different order from the rest; or (as already stated) the spiral
may change into the cyclical, or vice versa.

243. The relation of the phyllotaxy of a branch to the leaf
from the axil of which the branch springs is somewhat various.
But in Dicotyledons, the first leaf or the first pair of the branch is
mostly transverse ; that is, the first leaves of the branch stand to
the right or left of the subtending leaf. In Monocotyledons, the
first branch-leaf is usually parallel to and facing the subtending
leaf, as shown in Fig.'304.

244. When the internodes are considerably lengthened, the
normal superposition of leaves is not rarely obscured by torsion
of the axis: indeed, this may equally occur in short internodes,
sometimes irregularly or in opposite directions, sometimes uni-
formly in one. Thus, in Pandanus utilis, or Screw-Pine, of
tristichous arrangement, the three compact vertical ranks be-
come strongly spiral by a continuous torsion of the axis. The
later leaves of Baptisia perfoliata, which are normally distichous,
become one-ranked by an alternate twist, right and left, of the
successive internodes.

245. When the internodes are short, so that the leaves approx-
imate or overlap, it is difficult or impossible to trace the suc-
cession of the leaves on the primitive spiral, but it is easy to
see which are superposed. The particular phyllotaxy may then
be determined by counting the vertical ranks, which gives the
denominator of the fraction. But in compact arrangements
these vertical ranks are commonly less manifest than certain
oblique ranks, which are seen to wind round the axis in oppo-
site directions. (See Fig. 245, 246.) These are termed second-
ary spirals, also by some parastichies. These oblique spiral

1 That is, of the observer and as seen from without. See p. 51, foot-note.

128 PHYLLOTAXY, OR LEAF-ARRANGEMENT.

ranks are a necessary consequence of the regular ascending
arrangement of parts with equal intervals over the circumference
of the axis; and, if the leaves are numbered consecutively, their
numbers will necessarily stand in arithmetical progression on the
oblique ranks, and have certain obvious relations with the pri-
mary spiral which originates them, as will be seen by projecting
them on a vertical plane.

245°. Take, for example, the 2 arrangement, where, as in the
diagram annexed to Fig. 244, the primitive spiral, written on a
plane surface, appears in the numbers, 1, 2, 3, 4, 5, 6, and so
on: the vertical ranks thus formed are necessarily the numbers
1-6-11; 4-9-14; 2-7-12; 5-10-15; and 3-8-13. But two
parallel oblique ranks are equally apparent, viz. 1-3-5, which,
if we coil the diagram, will be continued into 7-9-11-15-15 ; and
also the 2—4—6-—8-—10 continues into 12-14, and so on, if the axis
be prolonged. Here the circumference is occupied by two secon- -
dary left-hand series, and we notice that the common difference
in the sequence of numbers is two; that is, the number of the
parallel secondary spirals is the same as the common difference of
the numbers on the leaves that compose them. Again, there are
other parallel secondary spiral ranks, three in number, which
ascend to the right ; viz. 1-4—7, continued into 10-15 ; 3-6-J-12,
continued into 15; and 5-8-11-14, &c.; where again the common
difference, 3, accords with the number of such ranks. This fixed
relation enables us to lay down the proper numbers on the leaves,
when they are too crowded for directly following their succes-
sion, and thus to ascertain the order of the primary spiral series
by noticing what numbers come to be superposed in the verti-
cal ranks. Thus, in the small cone of the American Larch
(Fig. 244), which usually completes only three heights of leaves,
the lowest, highest, and a middle one make a_ vertical row
which faces the observer. Marking this first scale 1, and count-
ing the parallel secondary spirals that wind to the left, we find
that two occupy the whole circumference. From 1, we number
on the scales of that spiral 3-5-7, and so on, adding the com-
mon difference 2, at each step. Again, counting from the base
the right-hand secondary spirals, we find three of them, and
therefore proceed to number the lowest one by adding this com-
mon difference, viz. 1-4—7-10; then, passing to the next, on
which the No. 3 has already been fixed, we carry on that se-
quence, 6-9, &c.; and on the third, where No. 45 is already
fixed, we continue the numbering, 8-11, &c. This gives us in
the vertical rank to which No. 1 belongs the sequence 1—6—11,
showing that the phyllotaxy is of the five-ranked, or 2 order.

DISTRIBUTION OF LEAVES ON THE STEM. 129

It is further noticeable that the smaller number of parallel sec-
ondary spirals, 2, agrees with the numerator of the fraction in
this the ? arrangement; and that this number, added to that of
the parallel secondary spirals which wind in the opposite direction,
viz. 3, gives the denominator of the fraction. This holds good
throughout ; so that we have only to count the number of par-
allel secondary spirals in the two directions, and assume the
smaller number as the numerator, and the sum of this and the
larger number as the denominator, of the fraction which ex-
presses the angular divergence sought. For this, we must, how-
ever, take the order of secondary spirals nearest the vertical
rank in each direction, when there are more than two, as in all
the higher forms. But, in all, it is necessary to count only the
most manifest secondary spiral of each direction in order to
lay down the proper number on the leaves or scales, and so deter-
mine the phyllotaxy.* Ina rosette of the leaves of Houseleek
(Fig. 245) and a cone of Pinus Strobus (Fig. 246), the num-
bers which can be seen at one view are appended, and in the
latter the conspicuous secondary spirals are indicated: one to
left with a common difference of 5; and two to the right, of
which the most depressed and prominent has the common dif-
ference of 3, the other, nearest the vertical, the common differ-
ence 8. The 14th leaf is superposed to the first, indicating the
ys; 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 plyllotaxy,
leads into interesting fields. But this sketch may suffice for
botanical uses.

246. Relations of Whorls to Spirals. Verticillate and alternate
phyllotaxy, or whorls and spirals, in all complete exemplifica-
tions, are to be considered morphologically as distinct modes,
not to be practically homologized into one. Nevertheless, transi-
tions between the two, and abrupt changes from one to the other
on the same axis are not uncommon, the former especially in
the foliage, the latter in the blossom. If the spiral be assumed
as the fundamental order, it is not difficult to form a clear con-
ception as to how such changes come to pass. <A single whorl

1 In applying this method to the determination of the phyllotaxy of a
cone, or any such assemblage of leaves, the student should be warned that,
although the cones of Pines and Firs are all normally on the alternate plan
(while those of Cypresses are on the verticillate), yet in individual cases
(common in Norway Spruce) the cone is plainly made up of pairs of oppo-
site scales which are spirally arranged. See note under 236.

130 PHYLLOTAXY, OR LEAF-ARRANGEMENT.

may most naturally be produced by the non-development of the
internodes between any two, three, or more alternate leaves.
Two proximate distichously alternate leaves would thus form a
pair; the three leaves belonging to one turn of the spiral in the
tristichous (4) arrangement would compose a trimerous whorl ;
the five leaves of the two turns in the pentastichous (?) arrange-
ment, a 5-merous whorl, &c. Verifications of this conception, by
whorls breaking up or reverting to spirals, are occasionally met
with, and the successive overlapping in spiral order of the
members of a trimerous or pentamerous whorl is very common.
The few instances among phzenogamous plants in which the
leaves are opposite and all in the same plane * (that is, the suc-
cessive pairs superposed) may be deduced from the distichous
alternate mode becoming opposite without further change, by
the simple suppression of alternate internodes. The frequent
disjunction of the members of the pair in similar and analogous
cases goes to confirm this view. But the characteristic of whorls
ordinarily is that proximate whorls alternate, that pairs de-
cussate. We cannot homologize this with spiral phyllotaxy ;
for in this lies the fundamental difference between the two plans.
We can explain it only by a reference to Hofmeister’s law, which
generally governs leaf-origination as to position, namely, that
succeeding leaves appear directly above the intervals between
the nearest preceding (241, note): this gives decussation or
alternation of successive pairs or whorls.*

247. Hypothesis of the origin of both. Instead of regarding
the spiral path on the stem which connects successive alternate
leaves as a purely formal representation, it may be conceived to
be the line along which the members in some original form were
physically connected, in the manner of a leaf-like expansion

1 As in Loranthus Europzus, &c., according to Braun. See 236, note.

2 This renders the verticillate an advantageous arrangement, perhaps no
less so than the distribution which spiral phyllotaxy effects. Both must be
considered to have been determined by and for their respective utilities, and
to have been independent determinations. For “there is no continuity or
principle of connection between spiral arrangements and whorls ” (Chauncey
Wright) ; since, although individual whorls are easily reducible to spirals,
each succession is an absolute break of that system.

As whorls of four members often (as especially in calyx, bracts, &c.) may
and sometimes should be viewed as two approximate pairs, so even the spiral
of five members, as in a quincuncial calyx, has been conceived to consist of
two whorls, one of two, the other of three leaves, the second alternating with
the first as nearly as possible. But this appears far-fetched and of loose
application. It is much clearer as well as simpler to regard the alternate
as the fundamental phyllotaxy, and to deduce individual whorls from spirals,
if need be, rather than to imagine spirals as somehow evolved from whorls.

DISTRIBUTION OF LEAVES ON THE STEM. 131

resembling a spiral stairway. Upon this supposition, the leaves
would be the relics, or rather the advantageous results, of the
segmentation of such a frond-like
expansion. the segments separatcd
through the development of the
stem in length and firmness, and
modified in the various adaptations
to the conditions of higher vege-
table life; even as leaves themselves
are modified into tendrils, bud-
scales, petals, or other usefully specialized structures. The type
on this conception would be a frond, consisting of

an elongating axis with a continuous leaf-blade on

one side, and this taking a spirally twisted form.

But the frond of Fucaceous Algze, Hepaticee, and the

like, is two-bladed. While a one-bladed frond, or

with one blade suppressed, might be the original of

alternate-leaved spirals, the two-bladed frond, simi-

larly broken up, would give rise to the opposite or

other varieties of verticillate arrangement.’

248. Fascicled Leaves need to be mentioned here,
in order that they may be excluded from phyllotaxy.
They are simply a cluster or tuft of leaves, belonging
to more than one node, and left in a crowded con-
dition because the internodes do not lengthen. They
may belong either to the alternate or the verticillate
series. In Barberry and in the Larch (Fig. 247),
they are evidently alternate ; and they may be inferred
to be so in Pines (Fig. 248), or even may be seen to
be so in the bud-scales which form the sheath sur- |
rounding the base of the 2, 3, or 5 foliage-leaves.
In Junipers, the leaves of the fascicles are in the
verticillate order.

1 This is the conception of the late Chauncey Wright. See his elaborate
and most suggestive essay in Mem. Amer. Acad. Arts and Sciences, ser. 2,
ix. 379, mainly reprinted in Philosophical Discussions (posthumous), 296-328,
in which the whole subject of phyllotaxy is acutely discussed, especially
in its relation to questions of origin and developed utilities. His conception

FIG. 247. Piece of a branch of the Larch, with two fascicles of leaves, 7. e. two very
short and stout branclilets, bearing scars of former leaves or bud-scales below, and a
dense cluster of leaves of the season at summit. The main axis bears scars from which
the alternate leaves of the developed axis of the preceding year separated.

FIG. 248. Piece of a branch of Pitch Pine, with three leaves in a fascicle or bundle
in the axil of a thin scale (a) which answers to a leaf of the main axis. The bunile is
surrounded at the base by a short sheath, formed of the delicate scales of the axillary
bud, of which the three leaves are the developed foliage.

132 PHYLLOTAXY, OR LEAF-ARRANGEMENT.

Section II. Disposition or LEAVES IN THE Bup.

249. Vernation and Estivation are terms in general use, under
which the disposition of leaves in the bud is treated. The first
relates to ordinary leaves in this early condition ; the second, to
the parts of a flower-bud ; not, however, as respects insertion,
or position on the axis, which is phyllotaxy (231), but as to

251 the ways in which they

249 20
are coiled, folded, over-
lapped, &c., either per se
yal bast or inter se. Prefoliation
and Prefloration are

etymologically better

terms, substituted by
6O Richard.?
250. The descriptive
= 253
=

terms which relate to
individual leaves or
parts, whether of foli-
age or blossom. mostly range themselves under the heads of
plications or of enrolling, and are such as the following, the
sectional diagrams of which are copied from the original figures

would make the two plans equally primordial. But the freedom with
which these actually interchange on the same axis greatly favors the less
hypothetical view that whorls may be condensed spirals. This assumes
only the well-known fact that internodes may be completely non-developed.

1 Better formed and more expressive terms: but the Linnean ones are
most in use, and, though fanciful, are not misleading. In English description,
it isas convenient and equally terse to say that the parts are imbricate, val-
vate, &., “in the bud.” Linnezus, in the Philosophia Botanica, described
these dispositions of leaves in the bud under the term Fo//atio, — not a happy
name,—but did not treat of them in the flower-bud. Later, in Termini
Botanici (Amen. Acad. vi. 1762, reprinted by Giseke in 1781), he intro-
duced the words Vernatio and ’stivatio in their now current botanical sense,
to designate, not the time of leafing and of flowering (spring and summer
condition), but the disposition of the parts in the leaf-bud and flower-bud
(at least of the petals) as respects foldings, coiling, &c., of single parts, and
modes of overlapping or otherwise of contiguous parts. The terminology
as regards single leaves, Linnzus fixed nearly as it now remains. That of
leaves or their homologues in connection, and as respects the flower-bud, was
very imperfectly developed until its importance (and much of its termi-
nology) was indicated by Robert Brown, in his memoir on Proteacez, 1809,
in the Prodromus a year later, and in other publications.

Ptyxis (the Greek name) is coming into use as a general term for the
folding, &c., of single parts. ;

FIG. 249-254. Linngan diagrams of sections of leaves in the bud. 249. Condupli-

cate. 250. Plicate or plaited 251. Convolute. 252. Revolute. 253. Involute, 254.
Circinate or Circinal j

VERNATION, OR PRAFOLIATION. 133

in the Philosophia Botanica of Linnzeus. They were applied
only to foliage, but they are equally applicable to floral parts.
Leaves, and all homologous or similar organs, if not simply
plane, will be either bent or folded or else more or less rolled up
in the bud. The first three of the following terms relate to the
former, the remaining terms to the latter. They are as to the
mode of packing

Plicate ov Plaited (Fig. 250), when folded on the several ribs,
in the manner of a closed fan, as in Maple and Currant. This
occurs only in certain palmately veined or nerved leaves.

Cenduplicate (Fig. 249), when folded lengthwise, or doubled
up flat on the midrib, as in Magnolia; a very common mode.
The upper face of the leaf is always within.

Reclinate or Inflexed, when the upper part is bent on the
lower, or the blade on the petiole, as in the Tulip-tree (the blade
of which is also conduplicate) .

Convolute (Fig. 251), when rolled up from one margin, 7. e.
one margin within the coil, the other without, as in Apricot
and Cherry.

Involute (Fig. 253), both margins rolled toward the midrib
on the upper face, as the leaves of Water Lily, Violet, c. ;
also the petals of Stcironema and Tremandra.

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

Circinal ov Circinate (Fig. 254), when coiled from the apex
downward, as the leaves of Drosera and the fronds of all the
true Ferns.

Oorrugate or Crumpled, as the petals of a Poppy, applies
to the irregular crumpling of the otherwise plane corolla-leayes.
This is a consequence of rapid growth in length and breadth
in a confined space.

251. The Ptyxis (or folding, &c.) of an individual leaf, of
which the foregoing modifications are the principal, should be
distinguished from the arrangement in the bud of the leaves of
a circle or spiral in respect to each other. The interest of the
latter centres in the flower-bud, 7.e. in wstivation. To this the
following exposition is devoted, although sometimes applicable
to leaf-buds also.

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

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

134 PHYLLOTAXY, OR LEAF—ARRANGEMENT.

their phyllotaxy, and partly of the way in which they comport
when their margins meet in growth. Those leaves which are
within, or of higher insertion on the axis, will almost necessarily
be enclosed or overlapped: those which are members strictly
of the same whorl or cycle may fail to come into contact, or
may meet without overlapping at the contiguous margins or
apex; yet they may be overlapped, since they may have grown
unequally or some a little earlier than their fellows. Conse-
quently, no perfectly clear line can be drawn in the flower between
cycles and spirals except by their mode of succession. More-
over, estivation strictly so called should be concerned only
with the disposition among themselves of the several members
of one whorl, or of one complete spiral. So the alternation of
contiguous whorls, as of the three inner with the three outer
flower-leaves of a Lily or a Tulip (the alternative sestivation of
DeCandolle), is a matter of phyllotaxy, not of wstivation. The
latter is properly concerned only with the relations of each three
leaves to each other.?

253. The proper estivations may be classified into those in
which the parts do not overlap, and those in which they do. Of
the first, there are two kinds, the open (est.
aperta) and the valvate, both characterized and
named by Brown.? Of the second, there is
one leading kind, the zmbricate (adopted by
Brown from Linnzeus), with subordinate modi-
fications. Accordingly, the vestivation is
said to be

254. Open or Indeterminate (est. aperta),
when the parts do not come into contact in the bud, so as to

1 The same applies to the two sets of sepals and of petals in Barberry, in
Menispermum, and of the petals in Poppy, &e. (359).

2 Linneus, indeed, has, “ stivatio valvata, si petala se expansura instar
glume graminis ponuntur,’—the name, but not the thing: the glumes of
grasses are not valvate in the botanical sense. So the term as to its proper
use may be said to originate with R. Brown.

8 For a brief discussion of “Estivation and its Terminology,” see Amer.
Jour. Sci. ser. 3, x. 339, 1875.

As to names, it is perhaps more correct to say of the cestivation that it is
imbricative, convolutive, valvidar, &e. (vest. imbricativa, convolutiva, valvaris, &e.),
but of the leaves or pieces, that they are imbricate, conrolute, valvate, &e., in
wstivation; but such precision of form will seldom be attended to in botan-
ical descriptions.

FIG. 255. Diagrammatic cross section of an unopened tlower of Linden: its outer
circle of floral leaves (sepals) valvate in the bud; the inner (petals) between convolute
and imbricate.

ZESTIVATION, OR PRAEFLORATION. 135

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

255. Valvate or Valvular, when the margins meet squarely in the
bud, without any overlapping, like the valves of a dehiscent cap-
sule. Familiar examples are aiforded by the calyx of the Linden
(Fig. 255) ; also that of the Mallow, Rhamnus, Fuchsia, and the
whole of the seyeral natural orders to which these belong. A
modification of this, caused by some induplication or involution
of the edges of the individual leaves, occurs in most species of

Clematis: in Clematis Virginiana, they Gf)
merely project within (valvate-indupli- (2 ot
cate); in Clematis Viticella, they are Dad 0

conspicuously involute (valvate-involute), ——

or yalvate with margins involute. Some- oa a
times (as in the calyx of certain Malvacew) the joined edges
project outwardly (or are valvate with reduplicate margins), but
only slightly so.

256. Imbricate or Imbricative is the general name for zstiva-
tion (or yernation) with overlapping. The name is taken from
the overlapping of tiles or shingles on a roof, so as to break
joints or cover edges. It was first applied, by Linnzeus, to
leaves or scales on a stem, when thickly set and incumbent in suc-
cessive ranks or heights, the upper partly covered by those next
below. ‘The involucre of an Aster or of the common Sunflower
is a typical illustration ; as also the leaves of a Camellia-fiower,
the sepals as well as the petals; and the sepals or outer leaves

of a Flax or a Geranium-flower afford a simpler but similar
instance, although, from the parts being nearly of the same size
and at the same height, the overlapping is lateral instead of
obviously from below. Fig. 258, 259, and the outer part of
260, also the inner leafy circle of 255, illustrate in diagram this
true and simple imbricative sestivation of a definite number of

FIG. 256. Valvate-induplicate flower-leaves (calyx) of Clematis Virginiana, &e.
257. Valvate-involute, asin C. Viticella.

FIG. 258-260. Imbricate zestivation: 258, in two whorls of three leaves each (calyx
and corolla); 260, same of five leaves in the outer circle, those of the inner circle con-
volute; 259, a single set of three imbricated leaves (in the corolla of Magnolia),
almost completely encircling each other.

136 PHYLLOTAXY, OR LEAF-ARRANGEMENT.

parts.!_ It is characteristic of it that some parts (one or more)
are wholly exterior or covering in the bud, and others (one at
least) interior or covered, at least the margins. Imbricative
eestivation, 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 the blossom are not
depressed spirals, but whorls or cycles (as may commonly be
the case in the corolla, but hardly 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 expressive of the particular grade of phyllotaxy (3, 4, 2,
3, &c.). But 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

Equitant, where leaves override, the older successively astride
the next younger. The typical instance is that of ancipital or
two-ranked (4) conduplicate leaves, successively clasping, at least
next the base, as in Iris, Fig. 217. In what Linnzeus termed
equitant-triquetrous (well seen in Fig. 240, 241), the leaves are
three-ranked (being of the 4 order), and each imperfectly
conduplicate.

Quincuncial sestivation (as in the outer part of Fig. 260) is
simply the imbricate estivation of five leaves (2), in which
necessarily the first and second are external, the fourth and fifth
internal, and the third with one margin external, where it over-
lies the fifth, and the other internal, where it is covered by
the first.

Alternative vestivation, as already stated (252), comes from
verticillate 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
Liliaceze), three members alternate with and are covered by the
other three, and this sort of imbricate sestivation is produced.
More properly, the two series are to be considered separately.
Where the parts are four (as in Fig. 395), the normal imbrica-
tion is decussate, two exterior and two interior. This is some-

1 All the examples referred to result from alternate or spiral phyllotaxy,
the former of higher series, the latter of the 4 (Fig. 258, 259), and of the 2
(Fig. 260) order. Instead of separating (with DeCandolle and others) the
2 arrangement as different in kind from»the imbricate (under the name of
quincuncial estivation), we should count it as a typical case. Otherwise the

¢ arrangement might equally claim a generic distinction, also the 3, &e.

ZESTIVATION, OR PRA FLORATION. 137

times a clear case of binary instead of quaternary, 7.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 portion of a
depressed spiral, say of the 2 order with one piece omitted, and
the others adjusted so as to fill the space.

257. There are various deviations from normally imbricative
estivation, 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
convolute sestivation, because the individual leaves are involute
in a manner approaching the convolute vernation of Linnzeus.
Another is the Vexillar, as in the Pea tribe (Fig. 306), where
members which should be external have somehow developed as
internal, both in calyxand 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-
strict the name of imbricative) is a state exactly intermediate
between the quincuncially imbricate and the convolute or

@ i) @

261 262 263
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 Convolute-imbri-
cate.’ To bring Fig. 261 back to the quincuncially imbricate

1 Tt would not be amiss, therefore, to name one of these modes, viz. that
of Fig. 261, Subimbricate, and the other, Fig. 262, Subconvolute. George

FIG. 261. Quincuncial imbricate mo lified toward convolute by one edge of the
second leaf developing inside instead of outside of the adjacent edge of the fourth.

FIG. 262. Convolute modified toward imbricate by one leaf having a margin inside
instead of outside its neighbor.

FIG. 263. Convolute, or convolutive, or contorted (twisted) «stivation, in diagram.
Tn these three diagrams, the dark circle above represents the position of the axis, the
flowers being axillary.

15 PHYLLOTAXY, OR LEAF-ARRANGEMENT.

form, we have only to reverse a single overlapping on the left-
hand side of the figure. To restore Fig. 262 to the convolute,
we have only to reverse a single overlapping at the lower right-
hand side. Changes like these, or the reverse, are not rare in
several species, particularly in the corolla. The normal form
and the deviation often occur in different flowers on the same
individual, thus indicating an easy passage between the imbricate
zestivation in the proper sense and the

258. Convolufe, otherwise called Obvolute or Contorted, or
Twisted, Fig. 263, and inner circle of Fig. 260. Here each leaf
successively overlaps a preceding and is overlapped by a following
one, all having a slight and equal obliquity of position, so that
all alike have an exterior and an interior (or a covering and a
covered) margin, and all appear to be as it were rolled up to-
gether. This is strikingly so when the parts are
broad and much overlapped, as in Fig. 264. Brown
( included this among the forms of imbricate zestiva-
( ey, tion, and so does Eichler, particularly distinguishing
St it, however, under the name here preferred. The

264 occasional transitions would justify such classifica-
tion. But in most cases it is so uniform, and in the corolla so
completely characteristic of whole families (such as Malvacez,
Onagracez, Apocynacez, Gentianaceze, Polemoniacez, &c.), and
is so distinct in its nature, that it may well take rank among
the primary kinds of zstivation. As to its nature, it is evident
that while the imbricate mode (at least the ternary, quinary.
&c.) indicates or imitates spiral phyllotaxy (some members be-
ing within or with higher insertion than others), the convolute
and the valvate (having all the members of the series on the
same plane) answer to verticillate phyllotaxy, or to whorls
instead of depressed spirals." The name which this mode of

i

if
(:

Henslow, in Trans. Linn. Soc. ser. 2, i. 178, proposes to call the former half-
imbricate; the latter (following the faulty French example) is his imbricate
proper.

The subimbricate mode has two varieties, distinguished by Eichler (in
Bliithendiagramme) as ascensive, when the lower or anterior (7. e. the pieces
next the subtending bract or leaf) are successively exterior (as in Fig. 261),
and descensive, when the covering is from the upper side, i. e. from the side
next the axis.

1 Still, as those members of a quincunx which normally should be wholly
external do sometimes become internal during their development in the bud,
similar changes may be conceived to change a quincuncial into a convolute
disposition ; but, to effect this, three out of the five overlappings would have
to be reversed.

FIG. 264. Convolute (also called contorted) wstivation of a corolla.

ZESTIVATION, OR PRAEFLORATION. 139

zestivation ought to bear is not yet well settled, but that of con-
volute, here preferred, will probably prevail.’

259. In recapitulation, these principal forms of the xstivation
of floral circles may be classified in a synopsis. They are: I.
those not closed = open or indelerminale: I. closed; and these

1, with the margins not overlapping = valeate ; 2, with margins
overlapping ; a, one or more with both margins covered = Bitbrs t-

cate; 6, all with one margin ‘covered, the other uncovered =
convolute.

260. Plicate or Plaited, when applied to the flower-bud as a
whole, is ina somewhat different category. The term is here
used for the plaiting

of a tube or cup,

composed of a circle

of leaves ‘eae
oT

into one body.

is well marked in
the corolla of as alpaind 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. 265); in the corolla of
most Gentians, the plaits are internal and straight.
In Conyolvulus and Datura (Fig. 266-268),
the narrow plaits overlap one another in a con-
volute way, when they are said to be Supercolute.
In the common Morning Glory and some other
species of Ipomoea, these plaits are besides spirally twisted or

1 See article entitled “ Hstivation and its Terminology,” above referred to.
The earliest name is Obvolute, given by Linneus to the kind of vernation in
which two leaves (conduplicate ones in his diagram) are put together so
that one half of each is exterior, the other interior. That is just the mode
in question reduced to a single pair of leaves, as it is in the calyx of a Poppy.
Mirbel is the only botanist who has applied the term to estivation, and to a
circle of more than two leaves, and it has never been adopted in botanical
descriptions. It has the disadvantage that the prefix o) to botanical terms
means obversely or inversely. Contorted (contorta), in English Twisted, is in
early and is the commonest use, and it is sometimes expressive. The objection
to it is, that contortion or twisting of the flower-bud often conspicuously oc-
curs where there is no overlapping of edges (as in many species of Ipomeea) ;
that really no twisting accompanies the overlapping in a majority of cases
of this xstivation; and that when there is a twisting it is not rarely in the
direction contrary to the overlapping; so that the contortion needs to be

FIG. 265. Cross section of the extrorsely plicate or plaited tube of the corolla of
a Campanula inthe bud. 266. Same of a Convolvulus (Calystegia), the plaits convolute
or supervolute.

FIG. 267. Upper part of unexpanded corolla of Datura; the plaits convolute or
supervolute. 268. Cross section of the same.

140 PHYLLOTAXY, OR LEAF—ARRANGEMENT.

contorted in the opposite direction; that is, the plaits overlap
to the right and are twisted to the left.’

261. Direction of Overlapping, &c. This is to be noted in the
ternary, quinary, or other forms of spirally disposed imbrication,
also in convolute and twisted or contorted estivation. It
may be either to the right (dextrorse) or to the left (sinistrorse).
The application of this term depends upon the assumed position
of the observer, whether outside or inside. We always suppose
him to stand outside, in front of the object: so when the over-
lapping is from right to left of the observer thus placed, as in
Fig. 266, it is sinistrorse; when from his left to right, as in
Fig. 267, 268, dextrorse.? The direction is generally constant,
but in many cases only prevalent, in the same plant or the same
species, or even the same genus: sometimes it is uniform or
nearly so throughout a whole natural order.

separately expressed. To describe the estivation in such cases as deztrorsum
contorta et sinistrorsum torta (or in similar English words), when the overlapping
is to the right and the twisting to the left, is at least awkward and cumbrous.
Convolute is a fitting name, of occasional early application to this estivation
(as by Jussieu to the petals of Malvaviscus), but without definition in this
sense ; it has for many years been steadily adopted by the present writer,
is employed by Eichler in Germany, and has recently been adopted by
G. Henslow and others in Great Britain. It has, however, the disadvantage
of having been used by Linnzus to express the coiling of single leaves, and
in a manner not wholly congruous, but still with one edge outside and the
other inside.

1 In our phraseology, dextrorsely convolute and sinistrorsely contorted ;
in the current phraseology above referred to, dextrorsely contorted or twisted
and sinistrorsely twisted!

2 The reasons for adopting this view (in opposition to the authority of
Linneus and DeCandolle) are given in note on p. 51.

ANTHOTAXY, OR INFLORESCENCE. 141

CHAPTER V.
ANTHOTAXY, OR INFLORESCENCE.

262. INFLORESCENCE, a term which would literally denote the
time of flower-bearing, was applied by Linnzeus to the mode, that
is, to the disposition of blossoms on the axis and as respects
their arrangement with regard to each other. ANTHOTAXY, a
name formed on the analogy of phyllotaxy, and denoting flower-
arrangement, is a better term. The subject really belongs to
ramification (83, 14-16), and is also concerned with foliation
and with phyllotaxy. It is most advantageously treated apart,
immediately preceding the study of the blossom itself.

263. In and near the _ blossom,
both axis and foliage very commonly
undergo modification, either abrupt or
gradual, giving rise in the former to
Peduneles and Pedicels, in the latter to
Bracts and Lractlets.

264. A Braet (in Latin Bractea) is a
leaf belonging to or subtending a
flower-cluster, or subtending a flower,
and differing from the ordinary leaves
in some respect, usually in shape and
size, not rarely in texture and color.’
They are commonly, but not always,
reduced or as if depauperate 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 often insignificant or 269
minute and functionless, sometimes obsolete (as in Cruciferae) , or

1 Bracts of the first order are sometimes called floral leaves (Folia floralia),
or at least these are not well distinguished from bracts. But the term floral
leaves is descriptively more properly and usually applied to leaves below
the bracts or proper origin of the flower-clusters, yet near them, and un-
like the proper cauline leaves. It is a vague term, and is in some danger of
being confounded (as it never should be) with another vague term, viz.
flower-leaves, or the leaf-like organs of the flower itself.

FIG. 269. Bract (spathe) of Indian Turnip, partly cut away below to show the
fleshy spike (spadix) of flowers wlich it surrounds.

142 ANTHOTAXY, OR INFLORESCENCE.

fugacious. Each flower is subtended by (grows from the axil
of) a bract in Fig. 277-280, &e. A cluster of flowers is sub-
tended by a conspicuous and colored bract in Fig. 269, 270, 271 ;
by a circle of colored bracts, imitating white petals, in Fig. 294.

Spatne is the name given to such an enclosing bract, or
to two or more leaves successively enclosing a flower-cluster.

Ixvo.ucre is the name given to a circle or spiral collection of
bracts around a flower-cluster, as in Cornel (Fig. 294, also in Fig.
280 and 286), or around a single flower, as in Hepatica and
Mallow. A compound inflorescence may have both a genera
and a partial involucre, one for the general flower-cluster, otheis
subtending the partial clusters. The name of involucre is then
reserved for the general one; that of

INVOLUCEL is applied to the partial, secondary, or ultimate
involucres.

Bractiets (Lat. Bracteola, diminutive of bract) are bracts of
e secondary or ultimate order. For example. in the slender
flower-cluster, Fig. 277, 6 is a bract, subtending each individual
flower-stalk ; b’ is a bractlet, or bract of secondary order, borne
on that partial flower-stalk itself. The French naturally translate
the Latin bracteola into bractéole (pl. bractéoles): in English,
bractlet is an idiomatic and better diminutive.

Patets (Lat. Palee), also called Chaff, are diminutive or

FIG. 270. Monophyllous spathe of Indian Turnip, with tip more erect. 271. Spathe
and spadix of Calla. 272. Raceme of Cherry, leafy at base. 273. Dichotomous cyme.
274. Panicle of Meadow-Grass, 275. A corymb.

BRACTS AND FLOWER-STALKS. 143

chaff-like bracts or bractlets on the axis (or receptacle) and
among the flowers of a dense inflorescence, such as a head of
Composite (275, Fig. 287, 288); and the name is also given
to an inner series of the

Guiumes of Grasses. These are peculiar chaffy bracts or bract-
lets which characterize the inflorescence of Grasses and Sedges.

265. Peduncle is the general name of a flower-stalk, that is,
of an axis or stem, which instead of foliage, or at least ordi-_
nary foliage, supports a
flower-cluster or a single
flower. In Fig. 276,
each peduncle (rising
from the axil of an ordi-
nary leaf, and therefore
answering to a branch)
bears a solitary flower. In Fig. 277, the peduncle bears a series
of flowers, or a flower-cluster... In this instance, each flower is
borne on a flower-stalk of its own, that is, upon a

PepiceL. This is the name given to distinguish
a partial flower-stalk, or, more strictly, the stalk
of each individual flower of an inflorescence. (Fig.
277-284.) In less simple flower-clusters, with
ramification of two, three, or more grades, general
peduncle, partial peduncles, and pedicels have to be
distinguished: the term pedicel is reserved for the
ultimate ramification.

Scape is the name given to a peduncle rising
from the ground, as that of most Primulas, of
Dodecatheon, Hepatica, and the so-called acaules-
cent or stemless Violets.

Ruacuis (backbone) is a name given to the axis
of inflorescence; that is, the continuation of the
stem or peduncle through a somewhat elongated
flower-cluster, as in a spike of Birch or of Plan-
tain, Fig. 289,290. When this axis is short, as in
a head (Fig. 285-288), it is usually called the
RECEPTACLE, a word also used for the axis or cauline- 277
part of a flower. The context should show when receptacle of
inflorescence, and when receptacle of the flower itself, is meant.
Both belong to axis or stem.

reo

¢
ae)
e

FIG. 276. Moneywort, Lysimachia nummularia, with axillary one-flowered
peduncles.

FIG. 277. A Raceme, with a general ee (p), pedicels (p’), bracts (b), and
bractlets (0).

144 ANTHOTAXY, OR INFLORESCENCE. °

266. Position of Flowers or Clusters. Flower-buds accord
with leaf-buds in origin, position, and structure, to this extent at
least, that the parts of both are leaves or homologues of leaves,
crowded in whorls or spirals upon a short portion of stem or
axis; aud as leaf-buds are either terminal or axillary (15, 75)
so also are flower-buds ; as a leaf-bud may give rise to a simple
or a compound growth, 7. e. may branch again and again, or not
branch at all, so flower-bearing branches, or the flower-bearing
extremity of a stem or branch, may bear a single flower, or a
more or less compound cluster. Thus, in Fig. 276, an axillary
peduncle, or naked branch, bears at apex a solitary flower; in
Tig. 277, a peduncle bears a loose cluster of flowers, each of
which springs from the axil of a small bract; in Fig. 285,
a terminal peduncle bears at summit a dense flower-cluster.
Flowers are either solitary or in clusters. When solitary, they
are naturally without bracts, being subtended instead (as in Fig.
276) by ordinary foliage.

267. ‘The elevation either of a solitary flower or a cluster on a
peduncle, or of individual flowers of a cluster on pedicels, is only
incidental. ‘The flowers may be stalkless, i. e. sessile.

268. The Kinds of Inflorescence which have received distinctive
names are various, but are all reducible to two types, which,
generally well marked, may sometimes pass into each other, and
which are not rarely combined in the same compound inflores-
cence.’ The two types differ in basis as do axillary from ter-
minal buds ; in the one the flowers are axillary or lateral, in the
other terminal in respect to the axis from which each flower or
its pedicel arises. But inasmuch as every flower, whatever its
position, is terminal to its own stalk or axis, it is better to dis-
tinguish the two types in other terms, and to name them the

269. Indefinite and Definite, or, in equivalent and similar terms,
the Indeterminate and Determinate.2 Each may be either simple
or compound. It is from the simple that the definitions are to
be drawn. In the former type, the rhachis or main axis of the
inflorescence is not terminated by a flower, but lateral axes, or
pedicels, are. In the latter, both the main or primary and the
lateral or secondary axes or stalks are so terminated. An inde-
terminate flower-cluster may go on to develop internode after

1 Inflorescence, as has been well insisted on by Guillaud (in Bull. Soe.
Bot. France, iv. 29), is a mode, not a thing. The things sometimes but in-
appropriately so called are flower-clusters, for which, if a general technical
name is needed, that of Anthemia, in English Anithemy, suggested by Guillaud,
is as good as any.

2 Also named by Eichler (Bliithendiagramme, 33, following Guillaud, 1. c.)
the Cymose and the Botryose type.

THE TWO TYPES. 145

internode of axis, and one or more leaves (bracts) at each node,
and then a flower in the axil of each bract, until its strength or
capability is exhausted. Or it may stop short with very few
flowers ; but the uppermost and youngest one will not really ter-

270 289

minate the rhachis (7. e. come from a terminal bud), though i:
may appear to do so. (Fig. 272, 277-279, &e.) The lower
flower-buds are evidently the oldest, and accordingly the first to
expand; and the expansion will proceed regularly from below
upward: wherefore this type of inflorescence has been called the
Ascending ov Acropeta!l ; likewise the Centripetal, because, when
the flowers are brought to the same level or near it (as in Fig.
279, 280) by a lengthening of the lower pedicels, with or with-
out relative shortening of the rhachis, the evolution
is seen to proceed from circumference to centre.
There is thus no lack of names; but, inasmuch as
the following type is commonly referred to under the
general name of Cymose, to this has recently been
given the counterpart name of Botryose. (271.)

281

270. A determinate flower-cluster (as seen in its gradual
development which is not rarely presented) has the last internode

FIG. 278-280. Diagrams of indefinite, indeterminate, contripetal, or botryose in-
florescence: 278, Raceme; 279, Corymb; 280, Umbel.

FIG. 281-284. Diagrams of definite, determinate, centrifugal, or cymose inflores-
cence: 281, a false or descending raceme; 282, a solitary terminal flower; 283, same with
two lateral flowers developing, forming a 3-flowered cyme; 284, same with lateral
peduncles 3-flowered, or a pair of 3-flowered cymes, beside the central or primary
terminal flower.

146 ANTHOTAXY, OR INFLORESCENCE.

of its axis terminated by a flower (Fig. 281-284), which answers
to a terminal bud. If more flowers appear, so as to compose a
cluster, they spring from the axils, preferably from the highest
axils, and are later. The order of evolution is shown in the figures
by the size of the flower-buds or degree of expansion of the
blossoms. Fig. 281 best shows why a determinate or definite in-
florescence is sometimes said to be Descending ; Fig. 283 shows
why it is called Centrifugal, the central flower first expanding ;
Fig. 284 exhibits the lateral or circumferential partial clusters
later than the central blossom, and their lateral flowers later
than their central.

271. Varieties of Indeterminate or Botryose Inflorescence. The
names of most of these have been fixed from the time of Linnzeus,
but defined without reference to the order of evolution of the
flowers. They are the Raceme, Corymb, and Umbel, with flowers
raised on pedicels; the Spike and Head, with sessile flowers ;
also some modifications of these, notably the Ament and the
Spadix. The raceme may be taken as the type. Sotrys is
equivalent to racemus, &c.; and, as the type includes diversity
of forms to which the name racemose would seem inapplicable,
the term botryose (botrytischen of Eichler) is best chosen as the
general name of it, and is a good counterpart to cymose for the
other type.

272. A Raceme (illustrated in Fig. 272, 277, and by diagram
in Fig. 278) is a simple flower-cluster, in which the flowers,
on their own lateral or axillary pedicels and of somewhat equal
length, are arranged along a relatively more or less elongated
rhachis or axis of inflorescence. The common Barberry, Cur-
rant, Choke-Cherry and Black Cherry, and Lily of the Valley
are familiar examples.

273. A Corymb (Fig. 275, 279) is a shorter and broader
botryose cluster, which differs from a raceme only in the relatively
shorter rhachis and longer lower pedicels; the cluster thus be-
coming flat-topped or convex. ‘The centripetal character is thus
made apparent. The greater number of the corymbs of Linnzeus
and succeeding botanists are cymes, the central flower first ex-
panding. And the term corymbose or corymb-like is still much
used in descriptive botany for a ramification which is mainly of
the cymose type, and where in strictness the term cymose should
be employed.

274. An Umbel (Fig. 280), as in Asclepias, &c., differs from
a corymb only in the extreme abbreviation of the rhachis or axis
of inflorescence, and the general equality of the pedicels which
thus all appear to originate from the apex of the peduncle, and

THE BOTRYOSE TYPE. 147

so resemble the rays of an umbrella; whence the name, and
whence also the pedicels or partial peduncles of an umbel are
termed its Rays. The bracts, brought by the non-development
of internodes into a depressed spiral or apparent (or sometimes
real) whorl, become an involucre. (264.) An umbel or any
similar cluster when sessile (without a common peduncle), and
the parts crowded, is sometimes called a Faseiele (or the pedicels
said to be fuscicled) ; but this term has been differently defined.
(280.) It is better not to use it for any special kind of inflores-
cence, but simply in the sense of a bundle of whatever sort.
This will accord with the sense in which it is applied to an
aggregation of leaves. (248.)

275. A Head or Capitulum (Fig. 285) is a globular cluster of
sessile flowers, like those of Red Clover, Button-bush, and
Plane-tree. The pedicels need not be absolutely wanting, but
only very short. An umbel ;
with pedicels much abbrevi-
ated thus passes into a head,
as in Eryngium, &c. And
ahead with rhachis elongated
passes into a spike. The
short rhachis of a head very
commonly takes the name of
receptacle. (265.) The whole
may be subtended by con-
spicuous bracts forming an
involucre (264) as in Fig.
286, or may be destitute of
any, as in Fig. 285. On ac-
count of the compactness
and mutual pressure under
growth, the bracts among ons
the flowers of such heads (normally one subtending each blos-
som) are apt to be rudimentary, reduced to little scales, or
abortive, or completely wanting. In the latter case, the recep-
tacle is said to be naked (nude), 7.e. naked of bracts: when
they are present, it is paleate or chaffy. A peculiar sort of head,
not undeserving a special name (though this is not necessary
in descriptive botany), is the

Antnopium, the so-called Compound Flower of the earlier
botanists, which gives the name to the vast order of Composite.

FIG. 285. Cephalanthus occidentalis, the Button-bush; a pair of leaves, and a
terminal peduncle bearing a dense head (capitulum) of flowers.

148 ANTHOTAXY, OR INFLORESCENCE.

The name means ‘‘ resembling a flower.” Although it has all
the characters of a true head, the resemblance to a flower is

remarkably striking, the involucre imitating a calyx, and the
strap-shaped (/igulate) corollas of the several flowers imitating
the petals of a single blossom. In some (such as Dandelion
and the Cichory, Fig. 286), all
the flowers of the head bear
these petal-like corollas; in
more (such as Aster, Sun-
flower, and Coreopsis, Fig.
287), only an outer circle of
flowers does so; the remain-
der, smaller and filling the
centre (or disk), may by the
casual observer be taken for
stamens and pistils, and further the deception. The rhachis
or receptacle of a head of this kind is commonly depressed,
bearing the flowers on what then becomes the upper surface,
which adds to the imitation.?

Syconium. This name, given to the Fig-fruit, should be here
referred to, as it is a sort of inflorescence, of the general nature
of a head, but with receptacle external and flowers enclosed

1 The receptacle of an Anthodium has been termed Clinanthium or Phor-
anthium ; and its involucre, a Periphoranthium or Periclinium. The head has
likewise been named a Cephalanthium.

FIG. 286. Flowering branch of Cichory, with two heads of ligulate flowers, of
natural size.
FIG. 287. Vertical section of a head of flowers of a Coreopsis.

THE BOTRYOSE TYPE.

149

within. See Fig. 657-659 (683), where its morpho.ogy is ex-
plained and illustrated. Viewed as an inflorescence, it has also

been named a HypantTHopium.

276. A Spike is a cluster of sessile (or apparently or nearly

sessile) lateral flowers on an elongated axis.

It may be de-

fined by comparison, as a head with the rhachis lengthened
(indeed a young head often becomes a spike when older), and

equally as a raceme with the pedicels
all much shortened or wanting. <A
common Mullein and a_ Plantain
(Plantago. Fig. 290) are familiar ex-
amples. Two modifications of the
spike (or sometimes of the head) gen-
erally bear distinct names, although
not distinguishable by exact and con-
stant characters, viz. : —

Spapix, a spike or head with a
fleshy or thickened rhachis. The
term is almost restricted to the Arum
family and Palms, and to cases in
which the inflorescence is accompanied
by the peculiar bract or bracts called
a spathe (Fig. 269-271). But the
two do not always go together: in
Acorus and Orontium there is properly
no spathe to the spadix; while in the
Iris family the bracts are said to form

290

a spathe, and there is nospadix. In Palms, the principal reason
for naming the inflorescence a spadix is its inclusion in a

spathe before anthesis.

FIG. 288. A slice of Fig. 287, more enlarged, with one tubular perfect flower (a) left
standing on the receptacle, and subtended by its bract or chaff (Ul); also one ligulate
and neutral ray-flower and part of another (c,¢): in d, d, the bracts or leaves of the

involucre are seen in section.

FIG. 289. Catkin of White Pirch. 290. Young spike of Plantago major.

150 ANTHOTAXY, OR INFLORESCENCE.

Ament or Catkry. This is merely that kind of spike with
scaly bracts borne by the Birch (Fig. 289), Poplar, Willow, and,
as to one sort of flowers, by the Oak, Walnut, and Hickory,
which are accordingly called amentaceous trees. Catkins usually
fall off in one piece, after flowering or fruiting. All true catkins
are unisexual.

277. Any of these forms of simple inflorescence may be com-
pounded. Racemes may themselves be disposed in racemes, spikes
in spikes (as in Triticum), heads
be aggregated in heads, umbels in
umbels, corymbs may be corym-
bosely compound, &e.; forming
compound racemcs, spikes, umbels,
and the like, the terminology of
which is easy. The most usual
case of truly homomorphous com-
pounding is that of umbels ; the
inflorescence of much the larger part of Umbelliferae being in
compound umbels, as in Fig. 290%. There is then the general
umbel, the rays of which become peduncles to
tue partial umbels, and the rays of the latter
are pedicels. Umbella and Umbellule desig-
nate in Latin terminology the general and its
partial umbels. Umbellets (coined by the late
Dr. Darlington) may well replace the latter
as the English diminutive. But umbels are
sometimes racemosely arranged, as in Aralia
spinosa, heads may be arranged in spikes,
and so on.

278. A Panicle, of the simple and normal
sort (as illustrated in Fig. 291), is produced
when a raceme becomes irregularly compound
by some (usually the lower) of its pedicels
developing into peduncles carrying several
flowers, or more than one, or branching again
and again in the same order. But in com-
pound clusters generally the secondary and
tertiary ramifications are apt to differ in type
as well as in particular mode, giving rise to

291 heteromorphous or mixed inflorescence. (288.)
As Linneeus defined the term, and as it has generally been em-
ployed in botanical descriptions, the panicle is a general term

FIG. 290¢ Compound umbel of Caraway. 291. A simple panicle.

THE CYMOSE TYPE. aaa

forvany loose and diversely branched cluster, with pedicellate
flowers. It is therefore difficult to restrict it in practice to the
indeterminate type. .

279. Varieties of Determinate or Cymose Inflorescence. The
plan of this type has been sufficiently explained. (270.) Its
simplest condition is that of a solitary terminal flower, peduncu-
late or pedicellate (as in Fig. 282), or sessile. The production
of more flowers, necessitates new axes from beneath, from the
axils of adjacent leaves or bracts. ‘These, being later, render
the evolution centrifugal. The simplest flower-cluster (unless
we call the solitary flower of Fig. 282 a one-flowered cluster) is
that of Fig. 283, where a secondary floral axis or peduncle has
developed from the axil of each leaf of the uppermost pair, or
where with alternate leaves there is a single uppermost leaf, and
then only one such peduncle, and thus is produced a three- (or
two-) flowered cymose cluster. The flower of the primary axis
is marked by its bractless peduncle (therefore a pedicel) ; the
lateral and secondary peduncles are known (commonly or nor-
mally) by their bracts or bract; the portion below the bracts
is proper peduncle; that above, of single internode, pedicel.
Bracts, like other leaves, have potential bud§ in their axils ; these
in an inflorescence give the third order of ramification, each
branch tipped with its flower; and so on.

280. The Cyme is the general name of this kind of flower-cluster
in its various forms. One of these very simple cymes, by itself or
as a part of a larger cyme, may be called a Cymule. The regular

292

cyme usually accompanies opposite or other grades of verticillate
leaves, but is not rare in the alternate arrangement. It is
readiest understood in an opposite-leaved plant with regular
opposite ramification, as in an Arenaria, Fig. 292. By its con-
stitution, a cyme proceeds from simple to compound. It mat-

FIG. 292. Dichotomous or biparous cyme (cyme bipare of Bravais, Dichasium of
Eichler) of Arenaria Michauxii.

152 ANTHOTAXY, OR INFLORESCENCE.

ters little whether its development is progressive, the flowers of
the ultimate ramifications expanding after the earlier have matured
fruit, and with subtending bracts conspicuous or foliaceous ; or
whether, as in Elder and Hydrangea (Fig. 293, and in Fig.
273), the bracts are minute and caducous or abortive, and the
ramification complete with all the flower-buds well formed before
the oldest expand, so that the whole is in blossom almost at the
same time. But a cyme may be properly said to be compound
when the primary axis in it is a peduncle instead of a pedicel,

293

and supports a cluster (cyme or cymule) instead of a solitary
central flower at the main divisions.t One form of the regular
cyme, on account of its compactness, is named the

GromeruLe. This is merely a cymose inflorescence, of any
sort, which is condensed into the form of a head, or approach-
ing it. Of this kind is the so-called head of Cornus florida, and
of the herbaceous C. Canadensis (Fig. 294), which shows the

1 The dichotomous or two-branched cyme is the commonest, but is some-
times marked by suppression of internodes; as, for example, where the
branches are apparently in fours, in an umbelliform way ; but these are two
sets of two, with the internode between the pairs extremely short; or where,
as in Elder, the branches or rays are five, in this case consisting of the same
two pairs and a central one, which is a many-flowered continuation of the
primary axis. Or 5-rayed cymes, &c., may be founded upon alternate leaves
with shortened internodes, the rays or peduncles axillary to them thus
brought into an apparent whorl.

Bravais distinguished cymes as mu/tiparos, with three or more lateral
axes ; biparous, with two, and uniparous, with only one (cyme multipare,
bipare, unipare). To these Eichler gives the substantive names, severally,
of Pleiochasium, Dichasium, and Monochasinm. Only the latter needs illustra-
tion ; the others being as it were compounds of this.

FIG. 293. Compound cyme of Hydrangea; with some neutral and enlarged mar-

ginal flowers.
“

THE CYMOSE TYPE. 158

composition best, on close examination. A condensed but less
capitate cyme, or cluster of cymes, was called by Raper and
DeCandolle a FascrcLe; and this terminology has been much
adopted. It is properly enough
said to be a fascicle, which, as
used by Linneus and others,
means a bundle, or close collection
of parts, whether leaves, pedun-
cles, or flowers; but a fascicte is
not necessarily a cyme (274), noris
there need of a special substantive
name for a compact cyme, which
may either be simply so called or
it may pass into the glomerule.
281. Botryoidal forms of Cymose
Type, or False Racemes, &c. The
regular cyme seldom continues
with all its ramifications. In
Fig. 292, after the second forking,
one of the two lateral peduncles
mostly fails to appear, and in some } a4 i
parts one of the bracts also; and ultimately the lateral peduncle
present is bractless, like the central, therefore equally incapable
of further ramification, being reduced to a pedicel of a single inter-
node. This suppression some-
times begins at the first fork-
ing or at the very base; and,
when followed throughout, it
reduces a biparous or dichoto-
mous cyme to one half, and,
converts this half (when the
axis straightens) into the sem-
blance of a raceme if the
flowers are pedicelled, or of a
spike when they are sessile. 296 27
Fig. 296 is a diagram of such an inflorescence as that of Fig. 292.
with one lateral branch uniformly suppressed at each division, the
wanting members indicated by short dotted lines. Cases exem-
plifying this occur in portions of the inflorescence of some of our

FIG. 294. Plant of Cornus Canadensis: flowering stem bearing a cluster of leaves
aboye, then continued into a peduncle, and terminated by a glomerule of very small
flowers; this subtended by a colored and corolla-like involucre of four bracts. 295. One
of the flowers taken from the glomerule, enlarged.

FIG. 296. Uniparous cyme or sympodial false raceme, with opposite leaves or bracts.

FIG, 297. Form of the same, with alternate leaves or bracts.

154 ANTHOTAXY, OR INFLORESCENCE.

smaller Hypericums, and notably in H. Sarothra, in which the
leaves are all reduced to bracts. It is not always easy to show
why this is nota true raceme. But the other bract of the pair, upon
that supposition, is unaccountably empty : the successive angular
divergence of each joint of the axis of inflorescence in the younger
part, which commonly runs into a coil, finds explanation in the view
that each portion is the lateral branch from the axil of the subtend-
ing leaf: and occasionally the other axil produces a similar one,
thus revealing the cymose character. When the bract from the
axil of which the missing branch should come disappears also, as
sometimes it does, and uniformly on the same side, a state of
things like that of the upper part of Fig. 297 occurs. The’same
figure may serve for the arrangement corresponding to that of
Fig. 296, only with alternate leaves. But then, close as the imi-
tation of a raceme here is, the position of each flower in respect
to the bract supplies a criterion. While in a true raceme the
flower stands in the axil of its bract, here it stands on the oppo-
site side of the axis, or at least is quite away from the axil.

282. Sympodial forms. The explanation is that the axis of
inflorescence in such cases, continuous as it appears to be, is
not a simple one, is not a monopode, but a synypode (110, 116,
notes), z.e. consists of a series of seemingly superposed inter-
nodes which belong to successive generations of axes: each axis
bears a pair of leaves (Fig. 296) or a single leaf (Fig. 297),
is continued beyond into a peduncle (or pedicel in these
instances), and is terminated by a flower. From the axil
promptly springs a new axis or branch, vigorous enough soon
to throw the adjacent pedicel and flower to one side: this
bears its leaf or pair of leaves, and is terminated like its prede-
cessor with a flower; and so on indefinitely. The fact that the
alternate leaves or bracts are thrown more or less strictly to one
side and the flowers to the other, in Fig. 297, shows that these
leaves do not belong to one and the same axis; for alternate
leaves are never one-ranked or disposed preponderatingly along
one side of an axis, as in this diagram, and as is seen in the
inflorescence of a Houseleek, &c.

283. A further difficulty in the morphology of clusters of this
class comes from the early abortion or complete suppression of
bracts. This is not unknown in botryose inflorescence, occurring
in the racemes, of almost all Cruciferee : it is very common in the
cymose of all varieties, and especially in the uniparous ones in
question, which characterize or abound in Borraginacexze, Hydro-
phyllaceze, and other natural orders. In some genera or species,
the bracts are present, or at least the lower ones; in others,

—

THE CYMOSE TYPE. 155

absent; in some, either occasionally present or wanting in the
same species or individual. It is only by analogy, therefore,
and by a comparison of allied plants, that the nature of some of
these flower-clusters can be made out. With the botanists of a
preceding generation, 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 qualification that they
are false racemes or spikes, or cymose racemes, and the like; or
else, by reversing the phrase, with stricter correctness they call
them racemiform or spiciform cymes, &e.

284. Commonly these false racemes or spikes (or botryoidal
cymes, if we so name them) are circinate or inrolled from the
apex when young, in the manner of a crosier, straightening as
they come into blossom or fruiting. _ Likening them to a scorpion
when coiled, the earlier botanists designated this as scorpzord.
As the coil is a helix, it has also been named helicoid.1 The
flowers are then thrown, more or less strictly, to the outer side
of the coiled rhachis, where there is room for them; and so these
false racemes or spikes are secund or unilateral. The particular
anthotaxy and phyllotaxy of the various sympodial.and botryoi-
dal forms of cymose inflorescence become rather difficult ; and
the sorts which have been elaborately classified into species
(and have no little morphological interest) are connected by
such transitions, and are based on such nice or sometimes theo-
retical particulars, that the terminology based on them is seldom
conveniently applicable to descriptive botany, at least as to sub-
stantive names. ;

285. One of the latest and simplest classifications of cymes is
that of Eichler in his Bliithendiagramme.?

1 Scorpioid and Helicoid have been carefully distinguished by later
morphologists, on account of some difference in the mode of evolution and
arrangement of the flowers along one side of the rhachis, by which they
become two-ranked in scorpioid, one-ranked in helicoid. But practically
the two kinds of clusters are not always readily discriminated; and in gen-
eral terminology a single name, with subordinate qualifying terms, is suffi-
cient. Scorpioid is the older and commoner one, therefore the most proper
to be used in the generic sense.

2 CyMOsE TYPE (classified without reference to bracts, which are so often
wanting) ; divided into

a. Lateral axes three or more: PLe1ocnasium, the multiparous cyme ot
Bravais.

B. Lateral axes two: Dicnasium, the Liparous cyme of Bravais.

y. Lateral axis one: Monocuasium, the uniparous cyme of Bravais.

The latter, or the corresponding divisions of the preceding sorts, may be
divided as follows:

156 ANTHOTAXY, OR INFLORESCENCE.

286. Sundry complications and obscurities are occasionally
encountered in anthotaxy or phyllotaxy, which cannot here be

x Lateral axes transverse to the relatively main axis.

1. Lateral axes in successive generations always falling on the same
side of the relatively main axis: ScuravuBet [screwlike] or Bostryx
[ringlet or curl], the uniparous helicoid cyme of Bravais.

2. Lateral axes falling alternately on opposite sides of the relatively main
axis: WicKEL or CINcINNUS [a curl], the uniparous scorpioid cyme of
Brayais.

* * Lateral axes medial [in the same plane] relative to the main axis.

3. Lateral axes in successive generations always on the back side of the
axis from which it springs: FXcneL, Ruiprpium [fan].

4. Lateral axes in successive generations always on the upper side of the
axis from which it springs: S1cneL, Drepanivm [sickle].

The subjoined simple diagrams from Eichler (Fig. 298-503) illustrate
these forms. The ramification is, given without the bracts, which theoreti-
cally or actually subtend the axes of each generation. The student may add
them, and so more readi:y apprehend the characters.

Eichler recognizes the forms with median (antero-posterior) position of
axes in Monocotyledons only.
It is natural to distichous phyl-
lotaxy, and it accords with
the general rule that, in mono-
cotyledonous plants, the first
leaf of the branch, or the com-
monly solitary bractlet of the
° peduncle, stands over against
70 and facing the bract or leaf
28 from the axil of which said
branch or peduncle springs, 7.e. is posterior and next the parent axis, as
shown in the diagram, Fig. 304, 305.

301 300 a be
: \
OQ-O-0-0-0~ OKs pp
ps = 0 ( 0)
a Oana

FIG. 298. Diagram of the Cincinnus. 299. Diagram of the Bostryx The flower-
axes numbered in succession.

FIG. 300. Diagram of the Rhipidium. 302. Ground plan, indicating the order of
evolution of the flowers.

FIG. 301. Diagram of the Drepanium. 303. Ground plan, the flowers evolved in
succession, from left to right.

FIG. 304. Diagram showing the position of bractlet or first leaf on a branch in
Monocotyledons: a is the primary, a the secondary axis; is bract, and b/ bractlet.

THE CYMOSE TYPE. VT

explained, except through full details: such as flowers standing
by the side of a leaf, or a small leaf by the side of' a larger one,

The transverse or oblique position of secondary axes or peduncles, as in
Fichler’s first two species, brings the flowers of the false raceme or spike
out of line of the sympodial axis and bracts, neither in the axils, as in true
racemes, nor opposite them, as in the Rhipidium and Drepanium, but on one
side of this plane or the other. This is most common in Dicotyledons (in
Drosera, Sedum, Sempervivum, and Hyoscyamus, in Borraginacee and
Hydrophyllacexw, &c.), and is not rare in Monocotyledons, especially with
tristichous phyllotaxy, as in Tradescantia. In the Bostryx, Fig. 299, the
bractlet is anterior or falls on the same side as the bract, or, in other terms,
the successive bracts are all on one side, the inner side, of the helix; and the
Drepanium (Fig. 301) is like it: this is the helicoid cyme of Bravais, &«.,
and its flowers are commonly one-ranked. In the Cincinnus or true scorpioid
_ eyme (Fig. 298), and equally in the Rhipidium (Fig. 300), the bracts fall
alternately on opposite or different sides of the sympodial rhachis, because
the single bract (b’) of each successive secondary axis (a’) stands next the
axis (a) and over against the bract (4) of the generation preceding. The
flowers in these generally fall into two parallel ranks (conspicuously so
when crowded) on the upper side of the rhachis, on which, in the cincinnus
or true scorpioid cluster, they are usually sessile or nearly so (or spicate),
as is well seen in Heliotropes, and in very many Borraginaceous and ‘Hy dro-
phyllaceous species, in Houseleek, Tradescantia erecta, &c. This comes
through antidromy, that is, the phyllotaxy of aC)
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 T
right alternately. Fig. 301 is a plan of this Ls
two-ranked unilateral arrangement. Whennot

too crowded, both Cincinnus and Rhipidium ‘ :

are apt to have a zigzag rhachis. ( IT ao.
These two last-mentioned kinds are so gen- a vie FA

erally alike in character, as are equally the ys

Bostryx and the Drepanium, that the four spe- i

cies may as well be reduced to two. As these 7. Ge ) =

severally include the scorpioid and the helicoid a ~ Vy, -

uniparous cymes of modern anthotaxy, these v Vo)

terms may be retained to designate them. Or,

if other terms in use be preferred to scorpioid Pd athe

and helicoid, the form with two-ranked flowers B

may be denominated Cincinnal, that with single- 2057

ranked Gostrychoidal. But in neither type is the rhachis always coiled
up, although commonly more or less so in the undeveloped state.

While these forms generally imitate racemes or spikes, it will be noted
that Fig. 300 specially imitates a corymb in form and in seeming acropetal
or centripetal evolution. And when, as in this figure, the bracts are all
absent, no obvious external difference remains.

FIG. 305%. Ground plan (from Eichler’s Bliithendiagramme, i. 38) of the scorpioid
inflorescence of Tradescantia erecta, between bract (B) below and axis (a) above:
I., IL., IIL, &c., the successive flowers: v' is the bractlet of the first and bract of the
second flower, and so the others in succession up to 7° and a small undeveloped one
beyond. The figure 1 affixed to each flower indicates the first floral leaf.

158 ANTHOTAXY, OR INFLORESCENCE.

or a pedicel or peduncle above and out of connection with the
leaf which should subtend and accompany it.’

287. Mixed Inflorescence is not uncommon. This name is
given to clusters or ramifications in which the two types are con-
joined. Being heteromorphous, they are almost necessarily com-
pound, the two types belonging to different orders of ramification.
But under it may be included cases of comparatively simple
inflorescence, at least in the beginning, some of which nearly
fuse the two types intoone. In the Teasel (Dipsacus), an appar-
ently simple head or short spike comes first into flower at the
middle, from which the flowering proceeds regularly to the base.
Had it begun at the top, it wovld answer to Fig. 281, which,
blossoming from above downward by simple uniflorous lateral
axes along a monopodial primary axis, is a simple racemiform
eyme, while it may also be called a reversed or determinate raceme.
Something of this sort may be seen in certain species of Cam-
panula, with virgate inflorescence, the terminal blossom earliest,
the others following irregularly, or partly downward and partly
upward. In C. rapunculoides, when rather depauperate and
the inflorescence simple, the evolution is that of a true raceme,
except that a flower at length terminates the axis and develops
earlier than the upper half of the raceme. In Liatris spicata
and its near relatives, the heads, on the virgate general axis,
come into flower in an almost regular descending order, or are
reversely spicate. If in Fig. 281 the lower pedicels were prolonged
to the level of the upper, a simple corymbiform cyme would be
seen, with simple centrifugal evolution, that is, regularly from
the centre to the margin; thisis the counterpart of the rhipidium
or fan-shaped cyme, of Fig. 300, in which the evolution of the
blossoms is as regularly centripetal. The explanation of the
paradox is not far to seek.

1 The position of a pedicel at the side of a bract in false racemes is ex-
plained in the foregoing note. It may occur in true racemose inflorescence
by the reduction of sessile secondary racemes down to an umbel of two
flowers, transverse to the bract (as in many species of Desmodium), and
thus seemingly lateral to it, or to a single flower on the right or left of it.
The coalescence of a pedicel to the axis for a considerable height above the
subtending bract in a simple inflorescence, or above the last leaf in a
sympodial one (concaulescence of Schimper), is common. So likewise bracts
or leaves may be for a good distance adnate to sympodial shoots, whether
peduncles or leafy flowerless branches. This (named _ recaulescence by
Schimper) is of most frequent occurrence in Solanacee (in Datura, Atropa,
most species of Solanum, &c.), and is tie explanation of their so-called
geminate leaves,-where a large leaf (reaily belonging lower down) has a
small leaf by the side of it. See Wydler in Bot. Zeit. ii. 689, &c., Sendtner
in Fl. Bras. x. 183, and Eichler, Bliithend. i. 199.

THE CYMOSE TYPE. 159

288. Compound mixed inflorescence is very various and com-
mon; but the combinations have hardly called for special terms,
being usually disposed of by a separate mention of the generat
and of the partial anthotaxy, or that of the main axis of inflo-
rescence and that of its ensuing ramification.' In Composite,
for instance, the flowers are always in true heads, of centripetal]
evolution. ‘The heads terminate main stems as well as lateral
branches, so that they are centrifugally or cymosely disposed.
The reverse occurs in all Labiatsze and most Scrophulariaceze,
where the flowers, when clustered, are in cymes, but these cymes
are from axils, and develop in centripetal order. It is this
arrangement which mainly characterizes the

Tuyrsus. A compound inflorescence of more or less elongated
shape, with the primary ramification centripetal or botryose, the
secondary or the ultimate centrifugal or cymose. To the defini-
tion is generally added, that: the middle primary branches are
longer than the upper and lower, rendering the whole cluster
narrower at top and bottom, and sometimes that it is compact ;
but these particulars belong only to typical examples, such as
the inflorescence of Lilac and Horsechestnut. In the former, the
thyrsus is usually compound. A loose thyrsus is a

Mrxep Panicre. It is seldom that a repeatedly branching
inflorescence of the paniculate mode is of one type in all its
successive ramifications. Either the primarily centripetal will
become centrifugal in the ultimate divisions, or the primarily
centrifugal will by suppression soon run into false botryose
forms, into apparent racemose or spicate subdivisions. So that
the name Panicle in terminology is generally applied to all such
mixed compound inflorescence, as well as to the homogeneously
botryose. (278.)

VERTICILLASTER is a name given to a pair of opposite and
sessile or somewhat sessile cymes of a thyrsus or thyrsiform
inflorescence, which, when full, seem to make a kind of verticil or
whorl around the stem, as in very many Labiate. The name
was originally given to each one of the pair of cymes ; but it is
better and more commonly used to denote the whole glomerule
or false whorl produced by the seeming confluence of the two
clusters into one which surrounds the stem.

1 Guillaud (in his memoir on Inflorescence, published in Bull. Soc. Bot.
France, iv.) proposes to designate as Cymo-otryes the mixed inflorescence
composed of cymes developed in botryose order, 7.e. the thyrsus; and
Botry-Cymes, the reverse case of racemes, &c., cymoscly aggregated. For
the former, the old name thyrsus serves appropriately and well.

160 ANTHOTAXY, GIR} INFLORESCENCE.

289. The Relations of Bract, Bractlet, and Flower should here
be noticed, although the subject in part belongs rather to the
section on Floral Symmetry. (515.)

290. Anterior and Posterior, otherwise called /nferior and
Superior, and therefore Lower and Upper,’ are primary relations
of position of an axillary flower with respect to subtending bract
and the axis to which the bract pertains. The flower is placed
between the two. The portion of the flower which faces the
subtending bract is the anterior, likewise called inferior or lower.

The opposite portion which faces the axis of

5 inflorescence is the posterior, or superior, or

upper. ‘The right and left sides are lateral.

~N (Fig. 304, 306.) These relations do not

v( f =) ee in a solitary flower terminating a
g simple stem; but when such an axis produces

©) axillary branches with a terminal flower, the

———4 relation of this flower to the preceding axis

Lm & and its leaf is manifest, just as in indetermi-
nate inflorescence.

291. Median and Transverse. The position of parts which lie
in antero-posterior line, or between bract and axis, is median.
Thus, in Fig. 504 and 305, the parts are all in the median plane :
in Fig. 306, the btactlets, b’, b’, are lateral or collateral, or (being
in the opposite plane) transverse.

292. Position of Bractlets. The rule has already been laid
down (285) that the first leaf of an alternate-leaved secondary
axis is in Monoctyledons usually median and posterior, that is,
farthest away from the subtending leaf (as in Fig. 304, 305) ;
in Dicotyledons, lateral or transverse. When these secondary
axes are one-flowered peduncles or pedicels, the leaf or leaves
(if any) they bear are bractlets.2 Commonly there is only this

1 Not (with propriety, although the terms have been so used) exterior or
outer for the anterior, and interior or inner for the posterior position. These
terms should be reserved for the relative position on the axis of successive
circles or parts of circles, spirals, &c. Covering or overlapping parts are
exterior or outer in respect to those overlapped.

2 Latin Bracteole: not that they are small bracts, but bracts of an ulti-
mate axis. In axillary inflorescence, the distinction between bractlet and
bract is obvious: in case of a solitary terminal flower, there is no ground of
difference: in terminal or cymose inflorescence, the difference is arbitrary ;
but we may restrict the term bractlet to the last bract or pair.

German botanists mostly distinguish between bracts, as a leaf subtending
a flower or cluster, and bractlets, by terming the former a Deckblatt, and the

FIG. 306. Diagrain (cross section) of papilionaceous flower and its relation to axis
(a), bract (b), and braetlets (b7, 67).

THE CYMOSE TYPE. 161

posterior one to a simple axis in Monocotyledons, and two
transverse ones in Dicotyledons, 7.e. one to the right and the
other to the left of the subtending bract, Fig. 306, 4’ b’.. When
the latter form a pair, they are perhaps always truly transverse ;
when alternate, they stand more or less on the opposite sides and
transverse. When more than one in Monocotyledons, they may
become either median or transverse, or even intermediate. The
relation of bractlets or bract, that is, of the last leaves of inflo-
rescence, to the first of the blossom, might be considered either
under Phyllotaxy or under Floral Symmetry. In general, it
may be noted that successive members stand over the widest
intervals ;! 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 bractiets, right and left, the first
flower-leaf is at } divergence from one (the uppermost) of them
when the circle is of three, or at 2 when of five members, or near
it; but with many exceptions.”

A tabular view of the kinds of inflorescence and their termi-
nology, serving as a key, may aid the student.®

latter, being the leaves which the new axis first bears, Vorb/éftter, which is
also the name they apply to primordial leaves in germination.

1 In accordance with Hofmeister’s law ; but (as Eichler remarks) not to
be explained on his mechanical principle of production in this place because
of the greater room: for the position of the first member of an axillary
flower is mostly the same as regards the subtending bract when the bractlets
are wanting.

2 When bractlets are wanting, the leaves of the first floral circle if two
are right and left; if three, two lateral-posterior and one anterior; when
five, the odd one commonly in the median line, either anterior or posterior.

3 INFLORESCENCE is either Pure, all of one type, or M1xep, of the two types
combined. The Types are:
I. Main axis not arrested and terminated by a flower. Indeterminate, Indefi-
nite, Acropetal or Ascending, Centripetal, or BorRYOSE.
II. Main and lateral axes arrested and terminated by a flower. Determinate,
Definite, Descending, Centrifugal, or Cymosr.

{62 ANTHOTAXY, OR INFLORESCENCE

I. BOTRYOSE TYPE.

1. Simple, with lateral axes unbranched and terminated by a single flower, and
Flowers on pedicels,
Of somewhat equal length on a comparatively elongated axis, RACEME.
The lower ones longer than the upper, and main axis short, . CoryMB.
Of nearly equal length on an undeveloped main axis, . . UMBEL.

Flowers sessile'‘on a very Short main axis, << . .. . . » BAD.
Flowers sessile on a comparatively elongated main axis,. . . SPIKE.
Acfleshy.spilke‘or head-is/ar Grice s34,. 1c.) 21a) ba) eS Ee
A sealy-bracted spike isan 5 - = 2 1 > a. = of = ae) SReANIIONIDIOR
CATKIN.

Compound, with lateral axes branched once or more, bearing clusters instead of
single flowers.
Irregularly racemosely or corymbosely compound,. . PANICLE.
Homogeneously and regularly compound, as

Racemes inaraceme, ... .. .. .. . ». ComMPpouND RACEME.

Corymbs: conymbose,..4). 2.) a) 5) Sh) . ) eComrounpiGonwane
Umbels inanumbel, . - .... 4.) .. « «5. 4. ComPouxDiUimern
Spikes spicate, . . 2. = «4c =. «+ » ~ COMPOUND) Spas

Homogeneously compound, the secondary ramification
unlike the primary, as Heads racemose, Umbels spiked,
Spikes panicled, &c.

If. CYMOSE TYPE.

1. Simple, with terminal axis of each generation one-flowered.
Monopodial, the axis of each oer as evidently re-
solved into branches, . . . . : 7 ew LERUE Onin
These more thantwo, . . . . iPicivehnetan or MuLTIPAROUS CYME.
These only two, . . .Dichasium, Dichotomous or BrrArous CYME.
Sympodial, the apparently simple axis continued by
a succession of new axes standing end to end,
Monochasium, False Raceme or Spike, Botryose or UNtpAROuUS CYME.
Flowers one-ranked on one side of rhachis, HELIcom Untrparous CyME.
Flowers two-ranked on one side of rhachis, Scorpiouy Unrparous CyME.

2. Compound, with terminal axes for one or more earlier
generations bearing a cyme instead of a single
flower, . .- . .- -. . . - Warious sorts of CompouND CYME.

Ill. MIXED INFLORESCENCE.

1. Anomalous simple, with unbranched one-flowered lateral
axes (287), suchas . . . PARTLY KEVERSED SPIKES OR RACEMES.
2. Compound, of various combirations, of which there are
names for the subjoined : —
Primary inflorescence botryose, with axis elongating;

secondary cymose, . . 5 ~ « « RYRSUR:
Pair of such opposite cymes yee caatuent round
thesmMaincaiShs phere een ey ws - « «© « « WERTICILLASTER:

Panicle with some of the ramifications ey mose, . . . MIXED PANICLE.

THE FLOWER. 163

CHAPTER VI.
THE FLOWER.

Section I. Irs Nature, Parts, AND METAMORPHY.

293. FLower-Bups are homologous with (morphologically an-
swering to) leaf-buds, and they occupy the same positions. (266.)
A FrLower is a simple axis or a terminal portion of one, ina
phznogamous plant, with its leaves developed in special forms,
and subservient to sexual reproduction instead of vegetation.

294. In passing from vegetation to reproduction, it is not
always easy to determine exactly where the flower begins. The
same axis which bears a flower or floral organs at summit bears
vegetative leaves or foliage below. Or when it does not, as
when an axillary flower-stalk or pedicel is bractless, the change
to actual organs of reproduction is seldom abrupt. Usually
there are floral envelopes, within and under the protection of
which in the bud the essential organs of the flower are formed.
Some or all of these protecting parts, in very many flowers, are
either obvious leaves or sufficiently foliaceous to suggest their
leafy nature; and even when the texture is delicate, and other
colors take the place of the sober green of vegetation, they are
still popularly said to be the leaves of the blossom. These pro-
tecting and often showy parts, though not themselves directly
subservient to reproduction, have always been accounted as
parts of the flower.1_ Between the lowest or outermost of these
and the bractlets and bracts there are various and sometimes
complete gradations. The axis itself occasionally undergoes
changes in such a way as to render the determination of the
actual beginning of the flower somewhat arbitrary. Moreover,
the flower itself is extremely various in different plants, in some
consisting of a great number of pieces, in others of few or only
one; in some the constituent pieces are separate, in others
combined. ‘The flower is best understood, therefore, by taking
some particular specimen or class of flowers as a representative

1 Indeed, the colored leaves, or envelopes in whatever form, essentially
were the flower in most of the ante-Linnean definitions (that of Ludwig
excepted), as they are still mainly so in popular apprehension.

164 THE FLOWER.

or pattern, and especially some one which is both complete and
morphologically simple.

295. Such a flower consists of two kinds of organs, viz. the
Protecting Organs, leaves of the blossom, or floral envelopes,
which, when of two sets, are Catyx and CoroLia; and the
Essential Reproductive Organs, which co-operate in the production
of seed, the STaMENs and PISsTILs.

296. Floral Envelopes, Perianth, or Perigone, the fioral leaves
or coverings. The former is a proper English designation of
these parts, taken collectively. But in descriptive botany,
where a single word is preferable, sometimes the name perianth
(Lat. perianthium), sometimes that of per/gone (or perigonium) ,
is used. Pertanthium,'! a Linnzean term, has been objected to,
because it etymologically denotes something around the flower ;
but it seems not inappropriate for the envelopes which surround
the essential part of the flower. Pertgonium,
a later term, has the advantage of meaning
something around the reproductive organs,
which is precisely what it is. Neither name
is much used, except where the perianth or
perigone is simple or in one set (when it is
almost always calyx), or where it is of two
circles having the gen-
eral appearance of one
and needing descrip-
tive treatment as such,
as in the petaloideous
Monocotyledons. Itis
also used where the
morphology is ambigu-
ous. Generally, the floral envelopes are treated distinctively as
calyx and corolla, one or the other of which (mostly the corolla)
may be wanting.

297. The Calyx is the outer set of floral envelopes. That is
its only definition. Commonly it is more herbaceous or foliaceous
than the corolla, and more persistent, yet sometimes, as in the
Poppy family, it is the more deciduous of the two. Not rarely it

1 Linneus (and about the same time Ludwig) used it in the sense of a
proper calyx, yet with some vagueness. Mirbel and Brown established it
in the sense of the collective floral covering. DeCandolle revived Ehrhart’s

FIG. 307. The complete flower of a Crassula. 308. Diagram of its cross-section in
the bud, showing the relative position of its parts The five pieces of the exterior
circle are sections of the sepals; the next, of the petals; the third, of the stamens
through their anthers; the innermost, of the five pistils.

FIG. 309 <A sepal; 310, a petal; 311, a stamen; and 312, a pistil from the flower
represented in Fig. 307.

ITS NATURE AND PARTS. 165

is as highly colored. A name being wanted for the individual
leaves which make up the calyx, analogous to that for corolla-
leaves, DeCandolle 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 Perars.!

299. ‘The floral envelopes are for the protection of the organs
within, in the bud or sometimes afterward. Also, some of them,
by their bright colors, their fragrance, and their saccharine or
other secretions, serve for allurement of insects to the blossom, to
mutual advantage. (504.) This furnishes a reason for neutral
flowers, those devoid of essential organs, which sometimes
occur along with less conspicuous perfect ones. ‘‘ The leaves of
the flower” are therefore indirectly subservient to reproduction.

300. The essential organs, being commonly plural
in number, sometimes need a collective name. Where-
fore, the aggregate stamens of a flower have been
called the ANDRactuM ; the pistils, the Gynasctum.?

301. The Stamens* are the male or fertilizing
organs of a flower. A complete stamen (Fig. 311,
313) consists of Firament (/), the stalk or support,
and ANTHER (a), a double sac or body of two cells, 313
side by side, filled with a powdery substance, Potten, which is
at length discharged, usually through a slit or cleft of each cell.

well-formed name of perigonium, and in the sense here given. But later (in
the Organographie) 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
botany are 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 is most convenient to treat the parts as forming one organ.

1 Fabius Columna, at the close of the sixteenth century, appears to have
introduced this term, or, as Tournefort declares, “ primus omnium quod
sciam Petali vocem proprie usurpavit, ut folia florum a foliis proprie dictis
distingueret.”

2 The male household and the female household respectively, terms in-
~ troduced by Reeper (Linnea, i. 437), in the form of andraceum and gyneeceum ;
but the diphthong in the latter should also be @ The orthography
andrecium and gynecium (early adopted by Bentham, in Labiatarum Gen. et
Spec.) is conformable to the Linnean Monacia, Diecia, &c.

3 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 Tournefort and later, to the filaments; and the anthers were termed

FIG. 313. Stamen, composed of /, filament, and a, anther, with cells opening
laterally and discharging pollen.

166 THE FLOWER.

302. The Pistils, one or more to the flower, are the female or
seed-bearing organs.’ A complete pistil is distinguished into
.e three parts: the Ovary (lati

Ovarium, Fig. 314, a, shown in verti-
seeete » cal section, and Fig. 315, by Linnzeus
named Germen), the hollow portion
at the base which contains the

come seeds ; the SryLe (4), or colum-
nar prolongation of the apex of the
ovary; and the Stigma (¢), a portion
of the surface of the style denuded of epidermis,
sometimes a mere point or a small knob at the
apex of the style, but often forming a single or
double line running down a part of its inner face,
and assuming a great diversity of appearance in
different plants. The ovary and the stigma are the
essential parts. The style (as also the filament of a
stamen) may be altogether wanting.

apices. It came in time to be used as now for the whole organ; but Lud-
wig (Inst. Reg. Veg.), in 1742, apparently first so defined it, and introduced
the term Anther for the Apex of Ray, or Theca of Grew.

1 Following Linnzus, this term is here freely used in the plural, and for
each actual separate member of the gynecium, each organ which has an
ovary, stigma, and commonly a style. Tournefort, who appears to have
introduced the word, employed it in the sense of gynecium. Many authors
define it thus, and then practically eliminate from botany this, one of the
oldest of its terms, and one by no means superfluous. The typical pistillum
of Tournefort is that of the Crown Imperial (Inst. i. 69, & tab. 1) and the
name is from the likeness to a pestle ina mortar. As it soon became im-
possible to apply the same name to the pistil of a Fritillaria or of a Plum,
the cluster of such organs in Caltha, and the capitate cluster and receptacle
of such organs in a Ranunculus or Anemone, Linnzus, and Ludwig before
him, took the idea of Tournefort’s name, and used it accordingly.

“ Pistillum est pars interior et media floris, que ex ovario et stylo com-
ponitur. . . . Ovarium est pars pistilli inferior, que futuri fructus delinea-
tionem sistit. . . . Stylus est pars pistilli ex ovario centro producta. .. .
Summitas styli vel ejus partium Stigma dicitur.” Ludwig, Inst. Reg. Veg.
41-45, 1742. Without mentioning the plural, the pistil is thus defined in a
way which necessitates its use. Linnzus (in Phil. Bot.) first defines Stamen
and Pistillum in the singular number, enumerating the three parts of the
latter, and afterwards (p. 57) declares that “ Pistilla differunt quoad

FIG. 314. Vertical section of a pistil, showing the interior of its ovary, a, to one
side of which are attached numerous ovules, d: above is the style, 6, tipped by the
stigma, c.

FIG. 315. A pistil of Crassula, like that of Fig. 312, but more magnified, and cut
across through the ovary, to show its cell, and the ovules it contains; also pulled open
below at the suture. At the summit of the style is seen a somewhat papillose portion,
destitute of epidermis, extending a little way down the inner face: this is the stigma.

OvuLes, or bodies destined to be--

ITS NATURE AND PARTS. 167

303. The Torus, or Receptacle of the flower, also named
Tuavamts,’ is the axis which bears all the other parts, that
upon which they are all dc
(mnediately or immediately) 3
inserted. These are all ho-
mologous with leaves. ‘This
is extremity of stem, or
floral axis, out of which the
organs described grow, in
succession, like leayes on
the stem; the calyx from
the very base, the petals
next within or above the calyx, then the stamens, finally the
pistils, which, whether several or only one, terminate or seem to
terminate the axis. (Fig. 316.)

304. Metamorphosis. If flower-buds are homologous with
leaf-buds, and the parts of the flower therefore answer to leaves
modified to special functions (293), then the kind of flower here
employed in explaining and naming these parts 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-
blanee, 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 petals, are homolo-

numerum,” etc., and so elsewhere, besides founding his orders on the num-
ber of pistils. Among even French authors, Mirbel (1815) writes, “Le
nombre des pistils n’est pas le méme dans toutes les espéces,” &c. Moquin-
Tandon freely refers to pistils in the plural, and Aug. St. Hilaire takes
wholly the view here adopted, distinguishing the solitary pistil into simple
and compound. DeCandolle, in Théorie Elémentaire, third edition, writes,
“Chaque carpel est un petit tout, un pistil entier, composé d’un ovaire, @un
style, et dun stigmate.” Of English authors, no other need be cited than
Robert Brown. The terms in question, then, are : —

Gynecium, the female system of a flower, taken as a whole.

Pistil, each separate member of the gynecium; this either simple or
compound.

Ovary, the ovuliferous portion of a pistil. Substituting a part for the
whole, this term is often used when the whole pistil is meant.

Carpel, or Carpid, or Carpophyll, each pistil-leaf ; whether distinct as in
simple or apocarpous pistils, or in combination of two or more to form a com-
pound or syncarpous pistil.

1 By Tournefort, and adopted by Ludwig. Receptaculum floris, Linnzus.
Thorus, Salisbury. Torus (the proper form), DeCandolle.

FIG. 316. Parts of the flower of a Stonecrop, Sedum ternatum, two of each sort,
and the receptacle, displayed: a, sepal; 6, petal; c, stamen; 4, pistil.

168 THE FLOWER.

gous with leaves; that the sepals are comparatively little, the
petals more, and the reproductive organs much modified from the
type, that is from the leaf of vegetation. This is simply what
is meant by the proposition that all these organs are transformed
or metamorphosed leaves. What would have been leaves, if
the development had gone on as a vegetative branch, have in
the blossom developed in other forms, adapted to other fune-
tions. Linnzeus expressed this idea, along with other more
speculative conceptions, dimly apprehended, by the phrase Vege-
table Metamorphosis. Not long afterwards, this fecund idea
of a common type, the leaf, of which the parts of the flower,
&c., were regarded as modifications, was more clearly and differ-
ently developed by a philosophical physiologist, Caspar Frederic
Wolff. Thirty years later, it was again and wholly independ-
ently developed by Goethe, in a long-neglected but now well-
known essay, on the Metamorphosis of Plants. Twenty-three
years afterwards, similar ideas were again independently pro-
pounded by DeCandolle, from a different theoretical point of
view ; and finally the investigation of phyllotaxy has completed
the evidence of the morphological unity of foliaceous and floral
organs.*

1 The contribution of Linnzus is on p. 301 of the Philosophia Botanica,
1751; and all that is pertinent is in the following propositions : —

Plumulam seminis sepius terminat aut flos aut gemma.

Principium florum et foliorum idem est.

Principium gemmarum et foliorum idem est.

Gemma constat foliorum rudimentis.

Perianthium sit ex connatis foliorum rudimentis.

His dissertation, Prolepsis Plantarum, in Ameen. Acad. vi. (1760), added
nothing but obscure speculations to the former comparatively clear
statements.

Kaspar Friedrich Wolff’s contribution is in his Theoria Generationis,
mainly concerning animals, published in 1759, and an enlarged and amended
edition in 1774. He first clearly conceives the plant as formed of two ele-
ments, stem and leaf, but develops only the morphology of the latter, and
under the hypothesis that leaves of vegetation become bud-scales or floral
organs, as the case may be, through degenerescence or diminution of vege-
tative force, which is renewed in the bud or in the seed.

Johann Wolfgang Géthe’s Versuch die Metamorphose der Pflanzen zu
erkliren was published in 1790, in 86 pages. For the translations and
reproductions, see Pritzel, Thesaurus. To the French translation by Soret,
with German text accompanying (Stuttgart, 1831), and also to that of Ch.
Martens ((Euv. Hist. Nat. de Geethe, Paris, 1837), are joined the author’s inter-
esting notes and anecdotes of later periods, down to 1831. The degenerescence
by diminution of vegetative force with renewals by generation, propounded
by Wolff, in Geethe’s essay takes the form of successive expansion and con-
traction of organs. ;

A. P. DeCandolle’s Théorie Elémentaire de Botanique appeared in 1813,

)
|

ITS METAMORPHY. 169

305. It will be understood that metamorphosis, as applied to
leaves and the like, is a figurative expression, adding nothing to
our knowledge nor to clearness of expression, but rather liable
to mislead. The substance of the doctrine is wnity 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 evidence may
be adduced, one from the normal, the other from teratological
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 from 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. Sepals, 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.

397. 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 by transitions or

a second edition 1519; a third (revised by Alphonse DeCandolle), in 1844,
is posthumous. ‘The Organographie Végétale, 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 DeCandolle,
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 DeCandolle’s father), and first
investigated by the late Karl Schimper and Alexander Braun, beginning
about the year 1829.

It is interesting to know that Wolff's work was wholly unknown to
Geethe in 1790, and that both Wolff’s and Geethe’s were unknown to DeCan-
dolle until after the publication of the second edition of the latter’s Theorie
Elémentaire, in 1819. When the Organographie appeared, the essay of Goethe
had come to light; and contemporary contributions to floral morphology by
Petit-Thouars, R. Brown, Dunal, and Reper, were adding their influence.

170 THE FLOWER.

intermediate forms. ,The gradual transition from ordinary foli-
age to bracts and bractlets is exceedingly common. In color
a oe and texture it is not rare to

i V oes, meet with bracts which vie
Wf oe with, or indeed surpass, pet-
- als themselves in delicacy and

> brightness ; ae in such cases

— : x aes oe for attraction.
Ee geazle Sage, Painted-Cup
en oe # (Castilleia) , and the Poin-
(Ae
Ae Bee settia, with other Euphorbias

us

, OSS of the conservatories, are ex-
i. = i \ 7 .

iXisx/ amples of this. In the flowers
~~ of Barberry, it is by a nearly

arbitrary selection that bractlets are distinguished from sepals ;.

in Calycanthus, in many kinds of Cactus, and in Nelumbium,
the same is true
as to bractlets, se-
pals, and petals; in
Water-Lily (Nym-
phea, Fig. 318),
there is a gradual
transition from the
sepals through the
petals to stamens ;
in Lilies and most
lily-like flowers, se-
pals are as brightly
colored as_ petals,
and commonly more
or less combined
318 with them. When
the perianth-leaves are of only one set, it is not at all by color or
texture that this perianth can be assigned to calyx or to corolla.
Normal transitions from a stamen to a pistil could not, in the
nature of the case, be expected.
308. Teratological Transitions and Changes. Teratology is
the study of monstrosities. These in the vegetable kingdom

FIG. 317. Cactus-flower (Mamillaria cespitosa), with bractlets, sepals, and petals
passing into each other.
FIG. 318. Series exhibiting transition from sepals to stamens in Nymphza odorata.

ITS METAMORPHY. wie

often elucidate the nature of organs.! The commonest of these
changes belong to what was termed by Geethe retrograde meta-
morphosis ; that is, to reversion from a higher to a lower form, as of
an organ proper to the summit 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 flower, better named in Latin flos
plenus. 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 supernumerary petals
increase in number; and
the various bodies that may
be often observed, inter- ,——~
z ie ~
mediate between perfect >.
stamens (if any remain) and \———~
the outer row of petals, — “—~~
from imperfect petals, with
a 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 the transformation 319
that has taken place. Carried a step farther, the pistils likewise
disappear, to be replaced by a rosette of petals, as in fully double

1 The leading treatises are Moquin-Tandon’s Tératologie Végetale, Paris,
1841, and Masters, Vegetable Teratology, London, published for the Ray
Society, 1869. An earlier publication deserves particular mention, viz. the
thesis De Antholysi Prodromus, by Dr. George Engelmann, Frankfort on
the Main, 1832.

2 To these abnormal changes, the term metamorphosis is obviously more
applicable; for here what evidently should be stamens, pistils, &c., on the
testimony of position and the whole economy of the blossom, actually ap-
pear in the form of some other organ: yet even here the change is only in
the nisus formativus; the organ was not first formed as a stamen, and then
transformed into a petal or leaf.

FIG. 319. A flower of the common White Clover reverting to a leafy branch; after
Turpin. Calyx with tube little changed, but lobes bearing leaflets. Pistil stalked;
the ovary open down the inner edge, and the margins of the pistil-leaf bearing leaves
instead of ovules.

172 THE FLOWER.

buttercups.! In these the green hue of the centre of the rosette
indicates a tendency to retrograde a step farther into sepals, or
into a cluster of green leaves. This takes place in certain blos-
soms of the Strawberry, the Rose, &c. Such production of

‘‘oreen roses,” and the like, has been appropriately called
chlorosis, ov by Masters chloranthy, from the change to green.

309. A monstrosity of the blossom of White Clover, long ago
figured by Turpin (Fig. 319), is such a case of foliaceous rever-
sion, in which even the ovules are implicated.

‘ ) The imperfect leaves which take the place of

ff) the latter may be compared with the leafy

/ tufts which form along the margins of a leaf

of Bryophyllum, by which the plant is often
propagated. (Fig. 322.)

310. The reversion of a simple pistil di-

rectly to a leaf is seen in the Double-flowering

Cherry of cultivation (Fig. 320, 521), usually

passing moreover, by prolification of the re-

; ceptacle, into a leafy branch.

ILO yA 311. The reversion of pistils to stamens is
rarer, but has been observed in a good number
of instances, in Chives, in the Horseradish,
in Gentians and Hyacinths, and in some Willows. In the
latter, the opposite transformation, of stamens to carpels, is
very common, and curious grades between the two are met with
almost every spring. So also in the common Houseleek, and
in perennial Larkspurs. Certain apple-trees are known, both
in the United States and Europe, in which, while the petals are
changed into the appearance of minute green sepals, the outer
stamens are converted into carpels, these supernumerary and
in the fruit superposed to the five normal carpels.? In Poppies,
many of the innermost stamens are occasionally transformed
into as many small and stalked simple pistils, surrounding the
base of the large compound one.

1 Jt must not be concluded that the supernumerary petals in all’such cases
are reverted stamens, or stamens and pistils. Some are instances of abnormal
pleiotaxy, 7.e. of the production of one or more additional ranks of petals
(better deserving the name of double flower), with or without reversion of
essential organs to flower-leaves.

2 These trees are popularly supposed to bear fruit without blossoming;
the reverted green petals being so inconspicuous that the flower is un-
noticed.

FIG. 320, 321. Green leaves from the centre of a blossom of Double-flowering Cherry,
one still showing, by its partial involution and its style-like apex, that it is a reverted
carpel, the other a small but well-formed leaf.

FIG. 322. Leaf or leaflet of Bryophyllum, developing plantlets along the margins.

ITS METAMORPHY. 133

312. Another line of teratological evidence is furnished by
prolification. The parts of the flower are, by the doctrine,
homologous with leaves, and no leaf ever terminates an axis.
Normally, in fact, the
axis is never prolonged
beyond the flower, but
abnormally it may be.
It may resume yegeta-
tive growth as a termi-
nal growing bud, either
from between the pistils
after the whole flower is
formed, or at an earlier
period, usurping the
central part of the
flower. Thus, when a
rose is borne on a pe-
duncle rising from the centre of a rose, which is
not very unusual, or a leafy stem from the top
of a pear (Fig. 323), the flower was probably
complete before the monstrous growth set in. In
Fig. 524, the reversion to foliaceous growth took
effect after the stamens but before the pistils
were formed. In rose-buds out of roses, the terminal proliferous
shoot takes at once the form of a peduncle; in the shoot from
the pear, that of a leafy stem.

313. Again, 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 a | eas
flower-bud) makes its appearance in the eee he!
axil of a petal or of a stamen; and it
may be clearly inferred that the organ

clusters of flowers, from the axil of petals of garden Pinks are
sometimes seen. <A long-pedunculate flower from the axil of a

FIG. 323. A monstrous pear, prolonged into a leafy branch ; from Bonnet.

FIG. 324. Retrograde metamorphosis of a flower of the Fraxinella of the gardens,
from Lindley’s Theory of Horticulture; an internode elongated just above the stamens,
and bearing a whorl of green leaves.

FIG. 325. A flower of False Bittersweet (Celastrus scandens) producing other
flowers in the axils of the petals; from Turpin.

174 THE FLOWER.

stamen of a species of Water-Lily (Nymphza Lotus) is figured
and described by Dr. Masters.!

5314. In the application of morphological ideas to the elucida-
tion of the flower, nothing should be assumed in regard to it
which has not its proper counterpart and exemplar in the leaves
and axis of vegetation.

Section II. FrLorat Symmetry.

315. The parts of a flower are symmetrically arranged around
its axis.”, Even when this symmetry is incomplete or imperfect, it
is still almost always discernible; and the particular numerical
plan of the blossom may be observed or ascertained in some of
the organs.

316. Adopting the doctrine that the parts of the flower are
homologous with leaves, the synimetry is a consequence of the
phyllotaxy. It is symmetry around an axis, not the bilateral
symmetry which prevails in the animal kingdom. For parts of
a flower disposed in a continuous spiral (which mostly occurs
when they are numerous), the arrangement is that of some order
of this kind of phyllotaxy, which distributes the parts equably
into superposed ranks. (237.) The much commoner case of

1 The fullest enumeration and discussion of the very various kinds of
abnormal structures and deviations in plants is to be found in the Teratology
of Dr. Masters, above referred to. Many technical terms are here brought
into use, which need not be here mentioned, except the following, which relate
directly to floral metamorphosis.

Phyllody (called Phy/lomorphy by Morren, “rondescence by Engelmann) is

the condition wherein true leaves are substituted for some other organs;

i.e., where other organs are metamorphosed into green leaves. There is
phyllody of pistils, ovules, filaments, anther, petals, sepals, &c.

Sepalody, where other organs assume the appearance of green sepals.

Petalody, where they assume the appearance of petals, as normally in
Pinckneya and Calycophyllum, in which one calyx-lobe enlarges and becomes
petal-like, and abnormally in Primroses where all the calyx-lobes imitate
lobes of the corolla (this has been termed Calycanthemy) ; also of the stamens
of common “ double flowers.”

Staminody, where other organs develop into stamens. Cases of this as
affecting pistils are referred to above: rarely sepals and petals are so affected.

Pistillody, where other organs develop into pistils, which most rarely
happens except with the stamens, as above mentioned.

2 Tt is stated that Correa de Serra (who published botanical and other
papers in London, Paris, and Philadelphia during the first twenty years of
the century, but who knew far more than he published) was the first botanist
to insist on the symmetry of the flower. It was first made prominent by De
Candolle, in the Théorie Elementaire, and elaborated in detail by A. St.
Hilaire in his Morphologie Végétale.

FLORAL SYMMETRY. ae

equal number of parts in a cycle, and the cycles alternating
with each other, is simply that of verticillate phyllotaxy. (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 ¢somerous.

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 symmetry 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 cycles alternate, as is the nature of true whorls.
But the superposition of successive parts is not incompatible
with symmetry of the blossom, although it is a departure from
the ordinary condition, assumed by botanists as the type. An
tsomerous 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 ; 7. e., with the members of each circle all alike in shape.

319. These two kinds of symmetry 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 phyto-
graphy, as above defined.

320. A Complete Flower is one which comprises all four or-
gans, viz. calyx, corolla, stamens, pistil.

1 This is not only the definition “ generally applied in English text-books,”
but that introduced by DeCandolle, adopted by St. Hilaire, and followed at
least by the French botanists generally. The innovating German definition,
of a recent date, is that 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 “ simply
symmetrical or monosymmetrical,” and those which can be symmetrically
divided by two or more planes, “doubly symmetrical or polysymmetrical,”
as the case may be. Now both these forms have a more expressive and
older terminology, adopted by Eichler, viz. : —

Zygomorphous, for flowers, or other structures, which can be bisected in
one plane, and only one, into similar halves (median zygomorphous, when
this is a median or antero-posterior plane, as it most commonly is; trans-
verse zygomorphous, 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 in two or more
planes into similar halves.

176 THE FLOWER.

321. Numerical ground-plan. Many flowers are numerically
indefinite in some or most of their kinds of members, as Ranun-
culus, Magnolia, and the Rose for stamens and pistils, Nym-
pheea for all but perLaps the sepals, many Cactacez for all but
the pistil, and Calycanthus for all four components. But more
commonly each flower is constructed upon a definite numerical
ground-plan ; and the number is usually low. Seldom, if ever,
is it reduced to unity in a hermaphrodite blossom (even Hip-
puris, with a single stamen and a single pistil, is not an un-
equivocal case), and probably never in a complete one. But there
are such extremely simplified flowers among those of a single sex.
In Monocotyledons, 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 Houseleeks.

322. To designate the particular plan, such familiar terms of
Latin derivation as binary, ternary, quaternary, quinary, senary,
&¢., are sometimes employed, denoting that the parts of the
flower are in twos, threes, fours, fives, or sixes. More technical
and precise terms, equivalent to these, are composed of the Greek
numerals prefixed to the word meaning parts or members, as

Monomerous, for the case of a flower of one member of each ;

LDimerous, of two, or on the plan of two members of each ;

Trimerous, of three, or on the plan of three members ;

Tetramerous, of four, or on the quaternary plan ;

Pentamerous, of five, or on the quinary plan ;

Hexamerous,' of six, or on the plan of six members to each
circle. But, in Monocotyledons, so-called hexamerous blossoms
are really trimerous, the sixes being double sets of three.

523. Pattern Flowers. These should be symmetrical, regular,

complete in all the parts and without ex-
; \)/ cess or complication of these, and with-

: * te Vig out any of the cohesions or adhesions
u " {/ C = \ which may obscure the type, or render it
3) &O less expressive of the idea that a flower
c W ee consists of a series of circles or spirals
= of modified leaves crowded on a short

my = axis. Wherefore the illustration Fig. 307,

with its diagram Fig. 308, may serve as a pattern pentamerous
or quinary flower; and Fig. 326, with its diagram, Fig. 327,

1 These may be shortly written 1-merous, 2-merous, 3-merous, and so on
to 10-merous (decamerous), 12-merous (dodecamerous), &c.

Fig. 326. Parts of a symmetrical trimerous flower (Tillea muscosa): @. calyx;
6 corolla; ¢c. stamens; d. pistils. 327. Diagram of the same.

FLORAL SYMMETRY. TUT

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, may be exhibited in such a
diagram of a pattern isostemonous
d-merous flower as that displayed

in Fig. 328.
ha 324. Diplostemonous Type. 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.t As _ the
petals alternate with
the sepals, so the first
series of stamens al-
ternates with the pet-
323 als, the second series 2
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

1 This view of the symmetry of the flower was first taken by Brown
(Obs. Pl. Oudney, in-Denham and Clapperton Tray. 1826, reprinted in Ray
Soe. ed. of Collected Works, i. 293). It is true that Brown declares the
same of the pistils; but that isnot made out. The evidence 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 very common
appearance in haplostemonous flowers among the Dicotyledons of vestiges
of a second series, or of bodies which may be so interpreted. The andreecium
or the blossom is said to be

Tsostemonous or Haplostemonous when the stamens are 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.

FIG. 328. Ideal plan of a plant, with the simple stem terminated by a symmetrical
pentamerous flower; the different sets of organs separated to some distance from each
other, to show the relative situation of the parts. One of each, namely, a, a sepal, ),
a petal, c, a stamen, and d, a pistil, also shown, enlarged.

FIG, 329. A pentamerous diplostemonous flower of Seduna.

178 THE FLOWER.

superposed to) the sepals, and the stamens of the inner series
stand before the petals; as in the diagram, Fig. 331.7
325. Flowers which completely
exemplify their type or symmetry
are rare, but most exhibit it more
or less. Each natural order or
group exhibits its own particular
floral type, or modification of the
common type.2 Some of these
modifications do not at all affect
the symmetry or obscure the plan
of the flower, except by combina-
tions which render the phylline
character of the floral envelopes and carpels
less apparent, such combinations being of
rare occurrence in foliage. Others gravely
interfere with floral symmetry, sometimes to
such degree that the true plan of the blossom
is to be ascertained only through extended
comparisons with the flowers of other plants
331 of the same order or tribe, or of related
orders. The symmetry of the blossom finds its explanation in
the laws which govern the arrangement of leaves on the axis ;
that is, in phyllotaxy. The deviations from symmetry and from
typical simplicity have to be explained, and in the first instance

1 For convenient reference and the avoidance of circumlocution, some
writers term the stamens which are before the petals ep/peta/ous, those before
the sepals episepalous ; but, as this prefix means upon, it is better to restrict
these terms to cases of adnation of stamens to these respective parts of the
_ perianth, and to distinguish as

Antipetulous, those stamens which stand before petals, whether adnate or
free, and

Antisepalous, those which stand before sepals. — These terms we find have

already been employed in this way by Dr. A. Dickson (in Seemann, Jour.
Bot. iv. 275), with the addition of a third, viz.

Parapeta/ous, for stamens which stand at each side of a petal, yet not
necessarily before a sepal, as in many Rosacez.

2 These particular types, with their modifications, are set forth in the
characters or distinguishing marks of the orders, tribes, genera, &e. The
best generally available illustrations of ordinal types are in Le Maout and
Decaisne’s Traité Général de Botanique, and in Hooker’s English edition
and revision, entitled A General System of Botany, Descriptive and Analyti-
cal, London, 1873. The best morphological presentation is in Eichler’s
Bliithendiagramme, &c. (Flower Diagrams, Constructed and Illustrated),
Leipzig, 1875.

FIG. 350. Opened flower of Trillium erectum. 331. Diagram of the same.

a

ITS GENERAL MODIFICATIONS. 179

to be classified. To have morphological value, such explanation
should be based upon just analogies in the foliage and other
organs of vegetation. Whatever is true of leaves and of the
vegetating axis as to position of parts, mode of origin and
growth, division, connection, and the like, may well be true of
homologous organs in the flower.

Section III. Varrous MoprricaTions OF THE FLOWER.
§ 1. ENUMERATION OF THE KINDs.

326. In the morphological study of flowers, these modifica-
tions are viewed as deviations from type. Their interpretation
forms no small part of the botanist’s work. They may be classed
under the following heads : —

1. Union of members of the same circle : COALESCENCE.

2. Union of contiguous parts of different circles: ADNATION.

3. Inequality in size, shape, or union of members of the same
circle: IrREGULARITY.

4. Non-appearance of some parts which are supposed in the
type: ABORTION or SUPPRESSION.

5. Non-alternation of the members of contiguous circles :
ANTEPOSITION Or SUPERPOSITION.

6. Increased number of organs, either of whole circles or
parts of circles: AUGMENTATION or MULTIPLICATION.

7. Outgrowths, mostly from the anterior or sometimes pos-
terior face of organs: Enation.

8. Unusual development of the torus or flower-axis.

9. To which may be appended morphological modifications,
some referable to these heads and some not so, which are in
special relation to the act of fertilization. These are specially
considered in Section IV.

327. These deviations from assumed pattern are seldom single ;
possibly all may coexist in the same blossom. Several of them
occur eyen in that one of the orders, the Crassulacez, which
most obviously exhibits the normal type throughout.

328. Thus, Sedum (Fig. 329), with two circles of stamens,
being taken as the true type (324), Crassula (Fig. 307) wants
the circle of stamens before the petals; Tilleea (Fig. 326) is the
same, but with the members symmetrically reduced from five to
three ; Rhodiola loses all the stamens by abortion in one half
the individuals and the pistils in the other, sterile rudiments
testifying to the abortion; Triactina has lost two of its five
carpels, and the three remaining coalesce into one body up to the

180 THE FLOWER.

middle; Penthorum (Fig. 335, 336) has its five carpels coales-
cent almost to the top, and usually loses its petals by abortion ;
in Grammanthes and Cotyledon (Fig. 332-334), the sepals are
coalescent into a cup and the petals into a deeper one, out of

which the stamens appear to arise, these being adnate to the
corolla. Symmetrical increase in’ the number of members of
each circle is no proper deviation from type, at least in this
family (in which flowers on the same plant sometimes vary from
5-merous to 4-merous and 6-merous) ; and in Sempervivum (to
which Houseleek belongs) these members are always more than
five and sometimes as many as twenty in each circle.

§ 2. REGULAR UNION OF SIMILAR PARTs.

329. Coalescence, or the cohesion by the contiguous margins
of parts of the same circle or constituent set of organs, is so fre-
quent that few flowers are completely free from it. The last
preceding figures show it in the gynecium and corolla. Fig.
471-476 further illustrate it in the corolla, and in various degrees
up to entire union; and Fig. 483-488 illustrate it in the andree-
cium. ‘he technical terms gwhich coalescence calls for, and
which are needful in botanical description, may be found under
the account of the particular organ, and in the Glossary. Such
growing together of contiguous members in the blossom is strictly
paralleled by connate-perfoliate leaves of ordinary foliage (212,
Fig. 215), where it more commonly occurs in upper leaves, and
in bracts, which are still nearer the flower.

330. It should now be hardly necessary to explain that the
terms coalescence, cohesion, union, and the corresponding phrases

FIG. 332. Flower of Grammanthes. 333. Flower of a Cotyledon. 334. The corolla
laid open showing the two rows of stamens inserted on it. 335. The tive pistils of
Penthorum, united. 336. A cross-section of the same.

UNION OF PARTS. 181

in the next paragraph, do not mean that the parts were once sepa-
rate and have since united. That is true only of certain cases. The
union is mostly congenital, equally so in the disks of foliage of
the Honeysuckle (Fig. 215) and in the corolla of a Convolvulus.
The lobes which answer to the tips of the constituent leaves of
the cup or tube are usually first to appear in the forming bud,
the undivided basal portion comes to view later. It might be
more correct to say that the several leaves concerned have not
isolated themselves as they grew. Accordingly, Dr. Masters
would substitute for coalescence and adnate the term inseparate.
But the common language of morphology needs no change, as it
consistently proceeds on the idea, and the prevalent fact, that
leaves are separate things, and that the tube, cup, or ‘* insepa-
rate” base of a calyx or corolla, consists of a cértain number of
these. It is no contradiction to this view that they developed
in union.’

§ 3. UNION OF DISSIMILAR OR SUCCESSIVE Parts.

331. Adnation is the most appropriate term to denote the organic
and congenital cohesion or consolidation of different circles, the

1 If it were *eriously proposed to change the language of descriptive
botany in this regard, consistency would require its total reconstruction, with
the abolition of all such terms as cleft, parted, &c.; for the structures in
question are no more cleft than they are united. While these convenient
and long-familiar terms are continued in use (as they surely will be), although
quite contrary to literal fact, it cannot be amiss to continue those, such as
connate, adnate, coalescent, &c., which imply and suggest the fundamental fact
in the structure of phenogamous and the higher cryptogamous plants, viz.
that leaves are normally unconnected organs. :

Whether fusion or separation is the more complex condition, and therefore
indicativé 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. Buta
phylogenetic view of the whole case may reverse this conclusion as respects
the blossom. ‘The course of development from thallus and frond to distinct
foliage on an axis, from little to full differentiation, is clearly a rise in
rank, as also is the differentiation of fdliage into ordinary leaves, petals,
stamens, and pistils. But there is as much differentiation in the flower
of a Convolvulus as of a Ranunculus, and more in that of a Salvia, a
Lobelia, and an Orchis. In all such flowers, the combination, the irregu-
larity, and the diversification in many cases of the members of the same circle,
all indicate complexity, greater specialization, and therefore higher rank.
The production of leaves distinct from the axis is one step in the ascending
scale: such specializations and combinations of these as occur in flowers are
higher steps ; and the most specialized, complex, and therefore highest in
rank are complete, corolliferous, irregular flowers, with a definite number of
members, and these combined in view of the adaptations by which the ends
of fertilization and fructification are best subserved.

182 THE FLOWER.

apparent growing of one part on or out of another, —as of
the corolla out of the calyx, the stamens out of the corolla, or
all of them out of the pistil. This disguises the real origin
of the floral organs from the receptacle or axis, in successive
series, one within or above the other. Organs in this condition
are also and rightly said to be Connate (born united) ; but, as
this term is equally applicable to the coalescence of members of
the same circle, the word Adnate is preferable, as applying to
the present case only. Adnation is heterogeneous organic co-
hesion or adhesion: coa-
lescence is homogeneous
cohesion or union.

332. Adnation occurs in
very various degrees, and
affects either some or all the
organs of the flower. Its
consideration introduces into
terminology several peculiar
terms, which may here be
defined in advance. Three
of them, introduced and
prominently employed by
Jussieu, depend upon the
degree of adnation, or the
absence of it, viz. :—

Hypogynous (literally be-
neath pistil) , applied to parts
which are inserted (7. e. are
borne) on the receptacle of
the flower, as in Fig. 336.
This is the absence of
adnation, or the condition
which corresponds with the
unmodified type.

Perigynous (around the
pistil) implies an adnation
which carries up the inser-
tion of parts (which always means apparent origin or place of
attachment) to some distance above or away from the recep-

20Q
538

FIG. 336%. Vertical section of a flower of the Common Flax, showing the normal or
hypogynous insertion of parts upon the torus or receptacle.

FIG. 237. Vertical section of a flower of the Cherry, to show the perigynous insertion,
or adnation to the calyx, of the petals and stamens.

FIG. 338. Similar section of the flower of the Purslane, showing an adnation of all
the parts with the lower half of the ovary.

UNION OF DISSIMILAR PARTS. 183

tacle, so commonly placing this insertion around instead of
beneath the pistil; whence the name. The perigyny may be,
as the figures show, merely
the adnation of petals and
stamens to calyx, the calyx
remaining hypogynous, as in
Fig. 337; or else the adna-
tion of the calyx, involving
the other organs, to the lower
part of the ovary, as in Fig.
3358, or up to the summit of
the ovary, while the petals
and stamens are adnate still further to the calyx, as in Fig. 339.
The latter passes into what is called

Epigynous (on the pistil), where the adnation is complete to
the very top of the ovary, and none beyond it, as in Fig. 340,
541. Yet here the parts so termed are not really on the ovary,
except where an epigynous disk (394) actually surmcunts it.

333. Adnation brings some other terms into use in botanical
descriptions, especially those of superior and inferior. In this
connection, these words (in Latin taking the form of superus and
inferus) denote the position in respect to each other of ovary and
floral envelopes, — not the morphological, but the apparent posi-
tion or place of origin. Thus, in Fig. 836 and in 337, the calyx
is ¢nferior, or in other words the ovary superior. Here real and
apparent origin agree, this being the normal condition, which
is otherwise expressed by saying that the parts are free, 7. e. free
from all adnation of one to the other. But, in Fig. 339-341, the

FIG. 339. Similar section of a flower of Hawthorn, showing complete adnation to
the summit of the ovary and of the other parts beyond.

FIG. 340. Vertical section of a Cranberry-flower, and 341, of flower of Aralia
nudicaulis, with so-called epigynous insertion of calyx, corolla, and stamens; the calyx
of the latter completely consolidated with the surface of the ovary, or its limb
obsolete.

154 THE FLOWER.

ovary is said to be inferior and the calyx superior, the calyx
and other parts, in consequence of the adnation of its lower part,
seeming to rise from the summit of the ovary.

334. Adnation of floral envelopes to pistil rarely extends
beyond the ovary; yet, in species of Iris having a tube to
the perianth, this tube is commonly adnate for most of its length
to the style. But when the calyx has its tube or portion with
united sepals prolonged, the petals and the stamens are usually
adnate more or less to it, 7.e. are inserted on the calyx. And,
when the petals are united and prolonged into a tube, the sta-
mens, being within the corolla, are commonly adnate to or
inserted upon this. .

335. No one doubts that the view is a true one which repre-
sents the perianth-tube as adnate to the style in Iris, petals and
stamens as adnate to calyx in the Cherry (Fig. 337), stamens
as adnate to base of corolla in Fig. 334, and a long way farther
in Phlox, &c. That the calyx is similarly adnate to the ovary
is nearly demonstrable in certain cases.

336. But, as the lower portion of a pear is undoubtedly recep-
tacle, or rather the enlarged 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 Cereus or other Cactacea has all the external char-
acters and development of a branch, so it is most probable that
in many cases the supposed calyx-tube adnate to an inferior
ovary is partly or wholly a hollowed receptacle (in the manner
of a Fig-fruit) ; that is, a cup-shaped or goblet-shaped develop-
ment of the base of the floral axis. This would bring the case
under § 7. (326, 495.)

§ 4. IRREGULARITY OF SIMILAR PARTS.

337. Irregularity, or inequality in form or in union of mem-
bers of a circle, is extremely common, either with or without
numerical symmetry. One or two examples may suffice.

338. Irregular flowers with symmetrical perfection, except in
the gyneecium, are well seen in the Pea Family, to which belongs
the kind of corolla called Papilionaceous, from some imagined
resemblance to a butterfly. (Fig. 342-344.) This flower is
5-merous throughout, has the full complement of stamens (10, or
two sets), but the gynecium reduced to a single simple pistil.
The striking irregularity is in the corolla, the petals of which
bear distinguishing names: the posterior and larger one, exter-
nal in the bud, is the Vexititum or Stanparp (Fig. 344, a);

INEQUALITY OF SIMILAR PARTS. 185

the two lateral next and under the standard, ALa& or Wines (4) ;
the two anterior, covered by the wings and partly cohering to

O-4

form a prow-shaped body (c), the Carma or Kret. The calyx
is slightly irregular by unequal union, the two upper sepals
united higher than the
other three. The sta-
mens are much more
coalescent,but with an
irregularity, nine com-
bined by the lower part
of their filaments, and
one (the posterior)
separate. (Fig. 345.) a

339. The plan and * stad
floral symmetry in the Locust-blossom and 4;
its relatives are little obscured by the irregu-
larities and the coalescence, hardly more so
than in the plainer flower of its relative,
Baptisia (Fig. 347, 348), in which the petals are somewhat
alike, and the ten stamens are distinct or unconnected. Only
the calyx is more irregular, by the union of the two posterior
sepals almost to the tip. (Fig. 348.)

548

FIG. 342. Diagram of flower of the Locust, Robinia Pseudacacia: a. axis of inflores-
cence; 6. bract; first circle of 5, calyx; five remaining pieces, corolla; next anthers,
10 in number; in the centre a single simple pistil. 343 Front view of Locust-flower
showing only the corolla. 344. This corolla displayed.

FIG. 345. Andreecium of the Locust, nine stamens coalescent, one distinct. 346
Same of a Lupine, all ten filaments coalescent below into a closed tube.

FIG. 3!7. Calyx and corolla of Baptisia australis. 348. Same with petals fallen,
showing ten distinct stamens and tip of the style.

186 THE FLOWER.

340. But in a Lupine-blossom, of equally near relationship,
a casual observer might fail to recognize the very same type,
although disguised only by cohesions. For while the two pos-
terior sepals are united to the tip on one side of the blossom, the
three others are similarly united into
one body on the anterior side, giving
the appearance of two sepals instead
of five: in the corolla, the two keel-
petals are more strictly united into a
slender scythe-shaped or sickle-shaped
body; so that the petals might with
the unwary pass for four: in the
andreecium, the coalescence includes all
ten stamens (Fig. 346), which is an
approach to regularity.

341. The 5-merous symmetry of the
Violet-blossom is complete until the
gyneecium is reached (but with only
one circle of stamens) ; the main irregu-
larity of the perianth is in the anterior
petal, with its nectariferious sac at base
(Fig. 349-351) ; the two stamens near-
est this send into the sac curious appendages, which the other
three do not possess; the gynecium is composed of three car-

3) pels coalescent into one compound ovary in a
manner hereafter explained. In Antirrhinum
yy and Linaria (Fig. 480, 481), there is a similar

UN
(j irregularity accompanying coalescence of the
petals, the anterior one being extended at base

Me WW into a nectariferous sac or hollow spur.! The
/ flower of a Lobelia (Fig. 488) has the same

See numerical plan and symmetry as that of Viola

351 (except that the gyncecium is dimerous) ; but

the members are adnate below and coalescent above, and the

corolla is irregular through unequal coalescence of the five petals,
and the absence of coalescence down one side.

1 PeLoria is a name given by Linnzus to an occasional monostrosity of
these flowers (imitated in sundry others), in which the base of every petal, or
answering part of the corolla, is prolonged downward into a sac or spur.
The sac is, morphologically considered, a departure from normal regularity :
in the monster, symmetrical regularity is restored by the development of four
more sacs.

FIG. 349. Flower of Viola sagittata. 350. Its sepals and petals displayed. 351.

Diagram of a Violet-blossom, from Eichler, with bract or subtenling leaf (below), a pair
of bractlets (lateral), and axis to which the subtending leaf belongs (above or posterior).

——

DISAPPEARANCE OF PARTS. 187

§ 5. DISAPPEARANCE OR OBLITERATION OF Parts.

342. Abortion or Suppression are somewhat synonymous terms
to denote the obliteration or rather non-appearance of organs
which belong to the plan of the blossom. Abortion is applied
particularly and more properly to partial obliteration, as where
a stamen is reduced to a naked filament, or to a mere rudiment
or vestige, answering to a stamen and occupying the place of
one, but incapable of performing its office ; suppression, to abso-
lute non-appearance. Such vestiges or abortive organs justify
the use of these terms, the more so as all gradations are some-
times met with between the perfect organ and the functionless
rudiment which occupies its place. Such obliterations, whether
partial or complete, may affect either a whole circle of organs
or merely some of its members. ‘The former interferes with the
completeness of a flower, and may obscure the normal order of
its parts. The latter directly interferes with the symmetry of
the blossom, and is commonly associated with irregularity.

343. Of parts of a Cirele. Among papilionaceous flowers
(338), different species of Erythrina have all the petals but one
(the vexillum, Fig. 344, a) much reduced in
size, in some concealed in the calyx, and in
every way to be ranked as abortive organs.
In Amorpha, of the same family, these four
petals are gone, leaving no trace, reducing
the corolla to a single petal. (Fig. 352, 355.)
This one is evidently the vexillum, both by
position and shape ; and the 5-merous type,
also the particular type of the family, are still
discernible in the five notches of the calyx,
the ten stamens, &c. In a related genus,
Parryella, even this last petal is wanting, and
the andreecium is straight, all irregularity thus
disappearing through suppression. om

344. Delphinium or Larkspur and Aconite or Monkshood
furnish good examples of flowers in which irregularity is aecom-
panied by more or less abortion. The calyx of the Larkspur
(Fig. 354-356) is irregular by reason of the dissimilarity of the
five sepals, one of which, the uppermost and largest, is pro-
longed posteriorly into a long and hollow spur. Within these,
and alternate with them as far as they go, are the petals, only

FIG. 352. Stamens and pistil of Amorpha fruticosa. 353. An entire flower of the
Same,

188 THE FLOWER.

four in number, and these of two shapes, the two upper ones
having long spurs which are received into the spur of the upper
sepal; the two lateral ones having a small but broad blade

raised on a stalk-like claw; and the place which the fifth and
lower petal should occupy (marked in the ground-plan, Fig. 356,

FIG. 354. Flower of a Larkspur. 355. The five sepals (outer circle) and the four
petals (inner circle) displayed. 356. Ground-plan of the calyx and corolla.

FIG. 357. Flower of an Aconite or Monkshood. 358. The five sepals and the two
small and curiously shaped petals displayed; also the stamens and pistils in the centre.
359. Ground-plan of the calyx and corolla; the dotted lines, as in Fig. 356, representing
the suppressed parts.

DISAPPEARANCE OF PARTS. 189

by a short dotted line) is vacant, this petal being suppressed,
thereby rendering the blossom unsymmetrical. In Aconite
(Fig. 857-359), the plan of the blossom is the same,
but the uppermost and largest of the five dissimilar
sepals forms a helmet-shaped or hood-like body ;
three of the petals are wanting altogether (their
places are shown by the dotted lines in the ground-
plan, Fig. 359) ; and the two upper ones, which ex-
tend under the hood, are so reduced in size and so
anomalous in shape that 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 by Linneeus
NEcTARIES, a somewhat misleading name, as_ they
are no more devoted to the secretion of nectar than
ordinary petals or other partsare. 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
gyneecium is 2-merous), but with corolla, and not rarely the
calyx, irregular through unequal union in what is called the bila-
biate or two-lipped manner. ‘The greater union is always median,
or anterior and posterior, and two of the coalescent members form
one lip, three the other. The two posterior petals form the
upper lip, the anterior and two lateral form the lower lip of the
corolla; in the calyx, when that is bilabiate, this is of course
reversed. In some, as in Sage and Snapdragon, the bilabiation
of the corolla is striking (Fig. 479-481), and readily comparable
to the two jaws of an animal; in others, the parts are almost regu-
lar. The suppression referred to is, in most of these cases, that
of the posterior of the five stamens, as in Fig. 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

1 These natural orders in which this occurs, or tends to occur, are the
Scrophulariacee (Snapdragon, Pentstemon, Mimulus, &c.), Orobanchacee
(Beech-drops), Lentibulacee (Bladderwort), Gesneracee (Gloxinia), Big-
noniacez (Trumpet Creeper, Catalpa), Pedaliacew (Martynia), Acanthacee,
Labiate (Salvia, Stachys), &c.

FIG. 360. A petal (nectary) of an Aconite, much enlarged.

190 THE FLOWER.

filament, —a monstrosity, indeed, but the monstrosity is here
a return to normal symmetry. The two stamens nearest the
suppressed or abortive one generally
share in the tendency to abortion, as
is shown by their lesser length or
smaller anthers: in the flower of
Catalpa, these two also are either im-
perfect or reduced to mere vestiges
(as in Fig. 563) : in very many other
plants of these families, even these
vestiges are not seen, and so the five
stamens are by abortion or complete
suppression reduced to two.

346. Suppression in the gyneecium
to a number less than the numerical.
plan of the flower (as shown in the
perianth) is of more common occur-
rence than the typical number, and
the reduction is comparatively con-
stant throughout the genus or order.
A papilionaceous or other leguminous
flower with more than one or with all
five pistils is exceedingly rare, and
except in one pentacarpellary genus
is a monstrosity. Suppression of
the interior is more common than
of exterior organs. Want of room
in the bud may partially explain
this.

347. Suppression of whole Circles. Such suppression or rather
non-production in the actual blossom of whole series of organs
which belong to the type, and indeed are sometimes present in
that blossom’s nearest relatives, is very common. It gives
occasion to several descriptive terms, which may be here defined
together. First, and in general, flowers are

Incomplete, in which any one or more of the four kinds of organs
is wanting, whatever these may be;

Apetalous, when the corolla or inner perianth is wanting ;

Monochlamydeous, where the perianth is simple instead of

FIG. 361. Corolla of Gerardia purpurea laid open, with the four stamens, the place
which the fifth should occupy indicated by a cross.

FIG. 362. Corolla of Pentstemon grandiflorus laid open, with its four stamens, and
a sterile filament in the place of the fifth stamen.

FIG, 363. Corolla of Catalpa laid open, with two perfect stamens and tlie vestiges
of three abortive ones.

DISAPPEARANCE OF PARTS. 191

double, in which case the wanting set is generally (but not quite
always) the inner, or the corolla ;

Dichlamydeous, when both circles of the perianth (calyx and
corolla) are present ;

Achlamydeous, when both are wanting, as in Fig. 365. (These
three terms are seldom employed. )

Unisexual (also Diclinous or Separated), when the suppression
is either of the stamens or the pistils. In contradistinction, a
flower which possesses both is Bisexual or Hermaphrodite.

Staminate, or Male, when the stamens are present and the
pistils absent ;

Pistillate, or Female, when the pistils are present and the
stamens absent ;

Monecious (of one household), when stamens and pistils oc-
cupy different flowers on the same plant ;

Diecious (of two households), when they occupy different
flowers on different plants ;

Polygamous, when the saine species bears both unisexual and
bisexual or hermaphrodite flowers. This may occur in various
ways, from the greater or less abortion of either sex, either on
the same or on separate individual plants ;— as Moneciously or
Dieciously Polygamous, according to the tendency to become either
monececious or dicecious. Recently Darwin has well distinguished
the case of

Gyno-diecious, where the flowers on separate individuals are
some hermaphrodite and some female, but none male only ; and
Andro-diecious, 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. (556, 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 astaminate one; a fertile flower, one which
is pistillate, if not also hermaphrodite.

348. Suppressed Perianth. Almost universally, when the peri-
anth is reduced to asingle circle, it is the inner, or corolla, which
is not produced. Or, rather, when there is only one circle or sort
of perianth-leaves, it is called calyx, whatever be the appearance,
texture, or color, unless it can somehow be shown that an outer
circle is suppressed. For since the calyx is frequently delicate
and petal-like (in botanical language, petaloid or colored, as in

192 THE FLOWER.

Clematis and Anemone, Fig. 364), and the corolla is sometimes
greenish or leaf-like, the only real difference between the two is
that the calyx represents the outer
and the corolla the inner series.
Even this distinction becomes arbi-
trary when the perianth consists of
three or four circles, or of a less
definite number of spirally arranged
members.

5349. Yet the only perianth obvi-
ously present may be corolla, as when

to the ovary and its border or lobes obsolete or wanting.’ Aralia
nudicaulis (Fig. 541) is an instance, likewise many Umbellifere,
some species of Fedia or Valerianella, the fertile flowers of Nyssa,
and those Compositze which have no pappus. For Pappus,
the name originally given to thistle-down and the like, answers
to the border or lobes of a calyx attenuated and depauperated
down to mere fibres, bristles, or hairs. The name is ex-
tended to other and less obliterated forms. (644, Fig. 631-633.)
When the obliteration is complete, as in Mayweed (Fig. 630),
in some species of Coreopsis, &c., the corolla seems to be simply
continuous with the apex of the ovary. A comparison with
related forms reveals the real state of things.?

350. So also in Hippuris, in which (along
with extreme numerical reduction of the other
floral circles) the calyx as well as corolla seems
to be wanting ; but the insertion of the stamen
on the ovary (epigynous) suggests an adnate
calyx, and near inspection detects its border.

351. Both calyx and corolla are really want-
ing in the otherwise complete and perfect (symmetrical and

1 In the flowers of the two common species of Prickly Ash (Zanthoxy-
lum) of the Atlantic United States, one has a double, the other a single
perianth (as shown in Gray, Gen. Illustr. ii. 148, t. 156): the position of the
stamens gives a presumption that the missing circle of the latter is the calyx ;
yet it may be otherwise explained. In Santalacee there are some grounds
for suspecting that the simple perianth, although opposite the stamens, is
corolla; and the foliaceous sepal-lobes of the female flowers of Buckleya
woud confirm this, if these are true sepals rather than adnate bracts.

2 In the pappus of Composite, every gradation is seen between undoubted
calyx, recognizable as such by structure as well as position, and diaphanous
scales, bristles, and mere hairs, wholly “ trichomes ” as to structure, although
in the place of “phyllomes ” and representing them.

FIG. 364. Flower of Anemone Pennsylvanica; apetalous, the calyx petaloi I.
FIG. 365. Achlamydeous flower of Lizard’s-tail (Saururus cernuus), magnided.

the calyx has its tube wholly adnate ~

2

DISAPPEARANCE OF PARTS. 193

trimerous) flowers of Saururus, Fig. 365. But achlamydeous
blossoms are usually still further reduced to a single sex.
352. Suppression of one circle of stamens is of very common
occurrence. It is seen in different species of Flax ; which have
mostly 5-merous perfectly symmetrical and complete flowers with
one set of stamens abortive. In some species (as in Fig. 367),

vestiges of the missing circle of stamens are conspicuous in
the form of abortive filaments, interposed between the perfect
stamens ; in others, these rudiments are inconspicuous or even
altogether wanting.

353. Suppressed Andrecium or Gynecium. This occurs with-
out or along with suppression in the perianth. In cases of the
former, vestiges of 370 _. oon
the aborted organs WW,
often remain to sig- ,
nify the exact nature
of the loss. Sepa-
ration of the sexes
(monecious, diceci-
ous, &c.) is the re-
sult of such suppres-
sion. In Menisper-
mum (Fig.368, 369),
this is accompanied
by an actual doub-
ling of both calyx
and corolla. ‘The
dicecious flowers of
Smilax are similarly
complete, except by
the abortion of one sex, but the calyx and corolla are single.

FIG. 366. Flower of a Linum or Flax. 367. Andreecium and gynecium; the former
of 5 perfect stamens, alternating with 5 rudiments of a second set.

FIG. 368, 369. Dicecious flowers of Moonseed, Menispermum Canadense: 368, Stami-
nate or male blossom; 369, Pistillate or female, but with six abortive stamens, before
as many petals.

FIG. 370. A catkin of staminate flowers of a Willow, Salix alba. 371. A single
staminate flower detached and enlarged (the bract turned from the eye). 372. A pistil-
late catkin of the same species. 373. A detached pistillate flower, magnified.

{94 THE FLOWER.

354. Combined with suppression of Perianth. This, which is
found in most amentaceous or catkin-bearing trees, in some with
partial suppression of perianth, is well illustrated
in Willows, the flowers of which are all achlamydeous
and diecivus. (347.) The little scale (gland or nec-
tary) at the inside of each blossom might be sup-
posed to represent a perianth, reduced to a single
piece ; but an extended comparison of forms refers
= it rather to the receptacle. Willow-blossoms (Fig.
370-372) are crowded in catkins, each one in the axil of a bract :
the staminate flowers consist of a few stamens merely, in this
species of only two, and the pistillate of a pistil merely. In
Salix purpurea, the male flower seems to be a single stamen
(Fig. 374); but it consists of two stamens, united into one
body. Here extreme
suppression is ac-
companied with co-
alescence of the
existing members.

359. Still more
simplified flowers,
but more difficult
to comprehend, are
those of Euphorbia,
or Spurge. These
are in fact moneeci-
ous; and the female
flower is a pistil, the
male is a stamen.
The pistillate flower
(of three carpels,
their ovaries united
into one three-lobed
compound ovary)
surmounts a slender peduncle which terminates each branch of the
flowering plant. (Fig. 375.) From around the base of this pe-
duncle rise other smaller and shorter peduncles, each from the
axil of a slender bract, and surmounted by a single stamen,
which represents a male flower. (Fig. 376, 377.) This umbel-like

FIG. 374. A separate staminate flower of Salix purpurea, with the stamens coa-
lescent (monadelphous and syngenesious), so as to appear like a single one.

FIG. 375. Flowering branch of Euphorbia corollata. 376. Calyx-like involucre
divided lengthwise, showing the staminate flowers around a pistillate flower (a). 377. A
more magnified staminate flower detached with its bract, a; its peduncle or pedicel 5,
surmounted by the solitary stamen,c. 378. Pistil in fruit, cut across, showing the
three one-seeded carpels of which it is composed.

SUPERPOSITION OF SUCCESSIVE PARTS. 199

flower-cluster 1s surrounded and at first enclosed by an involucre
in the form of a cup, which imitates a calyx ; and the lobes of this
cup (the free tips of the calyx-leaves) in the present species are
bright white, so that they exactly imitate petals. Here, then, is
a whole cluster of extremely simplified flowers, taking on the
guise of and practically behaving like a single flower, the invo-
lucre serving as calyx and corolla ; the one-stamened male flowers
collectively imitating the andreecium of a polyandrous blossom,
and surrounding a female flower which might pass for the pistil of
it. A series of related forms, from various parts of the world,
gives proof that.this interpretation is the true one.

356. Suppression of beth Andrecium and Gynecium. ‘This
occurs in what are termed Neutral Flowers (347), such as are
conspicuous at the margin of the cymes of Hydrangea (Fig. 295)
and of Viburnum lantanoides and Opulus, also at the margin of
the head of flowers of Sunflower, Coreopsis (Fig. 287, 288), and
the like. In these and most other instances, the perianth of
which only the flower consists is much larger and more showy
than in the accompanying perfect flowers: in fact, their whole
utility to the plant, so far as known, is in this conspicuousness.
No plant normally bears neutral flowers only ; but in cultivation
all sometimes become so by monstrosity, as in the form of Vibur-
num Opulus called Snowball or Guelder Rose, also in ‘ full
double” roses, pinks, &c. Occasionally flowers become sterile
and neutral by mere depauperation and abortion of perianth as well
as of essential organs, as in certain Grasses; but such are
mostly vestiges of flowers rather than neutral blossoms.

§ 6. INTERRUPTION OF NORMAL ALTERNATION.

357. Anteposition or Superposition is the opposition of succes-—
sive (or apparently successive) whorls which normally alternate.
This result is brought about in different ways, some of which are
obvious, while of some the explanation is hypothetical.

358. In the first place, there are cases of seeming anteposi-
tion, which are explained away on inspection. In a tulip, lily,
and the like, there is a perianth of six leaves and a stamen be-
fore each. The simple explanation is that the flower is not
6-merous, but 3-merous: there is a calyx of three sepals, colored
and mostly shaped like the three petals, which alternate with
these and are clearly anterior in the bud; next, three stamens
alternate with the petals or inner circle of the perianth; then
the three stamens of the inner circle, alternating with the preced-
ing, necessarily are opposite the three petals. as the first three are

196 THE FLOWER.

opposite the sepals. These organs altogether are in four whorls
of three, not in two of six members; and the pistil at the centre,
of three combined members, is the fifth and final whorl.

359. The Barberry family exhibits a similar seeming ante-
position, which is more striking on account of a multiplication
of the members of the perianth. The calyx is of six sepals in
two circles, the corolla of six petals in two circles, the stamens
equally six; and so each petal has a stamen before and a sepal
behind it. But, when properly viewed as a trimerous flower with
double circles of sepals and petals as well as of stamens, all is
symmetrical and normal. Menispermum in the related Moonseed
family is in the same case, but the flower is trimerous, as seen
in Fig. 369: in the male blossom this is obscured in the andree-
cium (Fig. 368) by a multiplication of the stamens.’ The
same thing occurs in the perianth and bracts of certain Clusiacee,
in which the members counted as in fours are superposed, and
in some of which the double dimerous arrangement with apparent
anteposition extends through the corolla; while, in other closely
related flowers, the corolla changes to simply tetramerous and to
alternation with the preceding four sepals. This passes, in the
same family and in the allied Ternstroeemiacez, into

360. Superposition by Spirals, as where five petals are ante-
posed to five sepals, by an evident continuation of pentastichous
phyllotaxy ; and the stamen-clusters of Gordonia Lasianthus
are probably in this way brought before the petals.” The flower
of Camellia is continuously on the spiral plan up to the gyne-
cium ; but upon one which, from the bracts onward, rises from the
4 to the 2 and 2 order or higher, throwing the petals of the rosette
in a full-double fiower into numerous more or less conspicuous
vertical ranks.

361. Anteposition in the Andrecium. It is in the andreecium
that real anteposition is most common, and also most difficult to
account for upon any one principle. Doubtless it comes to pass
in more than one way. This condition is chiefly noticed when
the stamens are definite in number, and mainly in isostemonous
and diplostemonous flowers. (524. )

362. With Isostemony. Vitis (Fig. 879-381), also Rhamnus
(Fig. 415, 416), and the whole Grape and Buckthorn families of

1 In Columbine (Aquilegia), multiplication of the stamens in successively
alternating 5-merous whorls similarly brings the andreecium into ten ranks;
so, when these stamens in double flowers are transformed into hollow-spurred
petals, these are set one into another in ten vertical ranks.

2 Gen. Illustr. ii. t.140. But the petals alternate with the sepals in the
ordinary manner of the flower, though their strong quincuncial imbrication
suggests the spiral arrangement.

e =
SUPERPOSITION OF SUCCESSIVE PARTS. 197

which they 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 = 8% 580
the receptacle, alternate with and inside
the stamens: there is no good reason to
suppose that they answer to a second
row of stamens. Allisostemonous Por-
tulacaceze have the stamens before the
petals ; and, when the stamens are fewer
than the petals, those which exist occupy
this position. Among the orders with
gamopetalous corolla, such anteposition
is universal in Plumbaginacez, Primu-
laceze, the related Myrsinaceze, and in
most Sapotaceze, in the latter usually
with some complications. a

363. The earliest and the most obvious explanation of the
anomaly is that of the suppression of an outer circle of stamens,
and to this view recent morphologists are returning.’ Observa-
tion supplies no vestige of proof of it in Rhamnacez and Vitacez ;
but, in the group of related orders to which the Primulacez be-
long, evidence is not wanting. For Samolus and Steironema
both exhibit a series of rudimentary organs exactly in the place
of the wanting circle of stamens, which may well be sterile fila-
ments. In the allied order Sapotacez, while Chrysophyllum
has in these respects just the structure of Primulacez, and
Sideroxylon that of Samolus, Isonandra Gutta (the Gutta-percha
plant) has a circle of well-formed stamens in place of the sterile
rudiments of the preceding; that is, alternate with the petals,

1 Fichler, Bliithendiagramme, passim, and in preface to Part II. xviii.,
relating chiefly to obdiplostemony. The principal opposing view is that of
St. Hilaire, Duchartre, &c., maintaining that corolla and stamens here repre-
sent one circle of organs doubled by median chorisis ; upon which see note
under a following paragraph. According to that hypothesis, there is no
andreecial circle in such blossoms, or only vestiges of one, but the petals have
supplied the deficiency by a supernumerary production of theirown! The
more plausible hypothesis of Braun, that of a suppressed interior circle
of extra petals, would restore the alternation, and make the extant sta-
mens the fourth floral circle, as does the adopted explanation. Braun’s
hypothesis, if it insists*that an extra row of petals is wanting, supposes the
suppression of that which very rarely exists; but, if of stamens, then the
supposed suppression is of that which is so generally present, or with indi-
cations of presence, as properly to be accounted a part of the floral type.

FIG. 379. Flower of the Grape Vine, casting its petals before expansion. 380. The

same, without the petals: both show the glands of the disk distinctly, within the
stamens. 381. Diagram of the flower.

.
198 THE FLOWER.

completing the symmetry of the blossom and the normal alterna-
tion of its members. ‘This explanation of the anteposition of a
single circle of stamens is the more readily received, because it
well accords with the idea here adopted, that the androecium
of a typical flower should consist of two circles of stamens.
(324.) ‘The only serious objections to this explanation rise out of
the difficulty of applying it to analogous anteposition when both
circles are present.

364. For Diplostemony, the condition of two circles of sta-
mens, each of the same number as the petals, is also itself very
commonly attended by anteposition. In normal or Direct
Diplostemony, — that which answers to the floral type com-
pletely, — the antisepalous stamens (324, note) are the outer
and the antipetalous the inner series, and the carpels when
isomerous alternate with the latter and oppose the sepals; the
alternation of whorls is therefore complete, as in the diagram,
Fig. 382. Such stamens, however, may actually occupy a single
line or coalesce into a
tube, without derange-
ment of the type. But
it as commonly occurs
that the antipetalous sta-
mens are more or less
exterior in insertion, and
then the carpels, when
isomerous, are alternate
with the inner and anti-
sepalous stamens, and therefore opposite the petals, as in the
diagram, Fig. 383. This arrangement takes the name of
Obdiplostemony. In it the normal alternation of successive
whorls is interrupted, so as to produce anteposition,

365. With Obdiplostemony. This condition prevails, more or
less evidently, in Ericacez, Geraniaceze, Zygophyllacez, Rutacez,
Saxifragacex, Crassulacee, Onagracez, &c. (but in some of these
with exceptions of direct diplostemony) ; also, accompanied
by a peculiar multiplication of members (380), in Malvacez,
Sterculiaceze, and Tiliacee. The explanation is difficult. The
hypotheses may be reduced to three, neither of which is quite
satisfactory. There is, first, the hypothesis of St. Hilaire, ap-
plied to this as to the preceding case (to Rhamnus, Vitis, &c.),
that these exterior antipetalous stamens belong to the corolline
whorl; in other words, that the petal and the stamen before it

FIG. 382. Diagram of pattern flower with direct diplostemony. 383. Diagram
of similar flower with obdiplostemony. Both from Eichler’s Bliithendiagramme.

*

SUPERPOSITION OF SUCCESSIVE PARTS. 199

(whether adnate to or free from it) answer to one leaf which
has developed into two organs by a deduplication (372) taking
place transversely. ‘This makes the inner,and ae
antisepalous stamens the third floral circle 7%, 2~ 7" <\
or the only truly andrecial one, and sym- / “Be \
metrically alternate with the petals on the one ( Se 2)
hand and the carpels on the other. The NY G 2 o}
second hypothesis conceives that there is a \\So ee oy
whorl suppressed between these antipetalous
stamens and the corolla : this, ideally restored,
gives symmetric succession and alternation to all the succeeding
whorls. The five glands in a Geranium-flower, alternate with and
next succeeding the petals (Fig. 354), were plausibly supposed to
represent this missing whorl, which according to Braun should be
an inner corolla; according to others rather a primary circle of
stamens. The third is the recent hypothesis of Celakowsky,
which Eichler adopts: this regards the antipetalous stamens as
really the inner or second circle, and conceives that in the course
of development it has become external by displacement. The
difficulties of this hypothesis are, first to account for this dis-
placement, and then for the anteposition of the carpels to the
assumed inner stamens in the great majority of these cases.?

384

1 Jn the first part of the Bliithendiagramme, Eichler inclined to the first
hypothesis, that of St. Hilaire (now very much abandoned on account of
the feeble evidence that there is any such thing as transverse or median
chorisis); in the second, he discards this in favor of Celakowsky’s view
(published in Regensburg Flora, 1875). As to members which are morpho-
logically interior becoming exterior by outward displacement, Eichler cites
the staminodia or sterile stamen-clusters of Parnassia (Fig. 400, 401), and
the corresponding antipetalous stamens of Limnanthes, as clearly interior
in the early flower-bud, but exterior at a later period ; states that the vascu-
lar bundles which enter these stamens generally are either inner as respects
those of the episepalous stamens or in line with them; that in some cases
(as in many Caryophyilacee) the real insertion of the stamens is that of
direct diplostemony, while the upper part of their filaments and the anthers
are external to the episepalous series; that in most families with obdiplos-
temony examples of direct diplostemony occur, and still more cases with both
stamineal circles inserted in the same line; and that, as a rule, the episep-
alous stamens are either later or not earlier formed than the epipetalous.
As to the position of the carpels before antipetalous stamens and petals,
Celakowsky suggests that this may result from the outward recession of
those stamens affording more room there, while in the normal case the
greater space is over the episepalous stamens. And, indeed, exceptions
to the prevalent position are not uncommon both in direct diplostemony

FIG. 384. Diagram (cross-section) of the flower of Geranium maculatum, exhibiting
the relative position of parts, and the symmetrical alternation of circles, @. e. sepals,
petals, greenish bodies called glands, antipetalous stamens, antisepalous stamens,
carpels.

200 THE FLOWER.

366. The case of stamens in a cluster before the petals is a
complication of either of the foregoing with a peculiar kind of
multiplication, termed deduplication or chorisis. (372.)

§ 7. INCREASED NUMBER OF Parts.

367. Augmentation in the number of floral members is one
of the commonest modilications of the type. It occurs in two
ways: 1st, by an increased number of circles or turns of spirals
in the flower, which is Regular Multiplication ; 2d, by the pro-
duction of two or three or of many organs in the normal place
of one, Chorisis or Deduplication. ‘The first does not alter the
normal symmetry of the blossom, although it may render it dif-
ficult or impossible to trace or demonstrate it. The second
apparently disturbs, or at least disguises, floral symmetry.
Either may be definite, or of a constant and comparatively
small number; or c¢ndefinite, when too numerous for ready
counting, or inconstant, as the higher numbers are apt to be.

368. Regular Multiplication, or Augmentation of flural circles
or spirals, may affect any or all the four organs, but most com-
monly the andreecium. When the perianth is much increased
in the number of its members, the distinction between calyx and
corolla, or even between bracts and corolla, is apt to disappear,
as in most Cactaceous flowers (Fig. 317), Nelumbium, Calycan-
thus, &c. In these and similar cases, the members of the perianth
are prone to take a spiral instead of cyclic arrangement ; and this

and in obdiplostemony. Along with the lack of clear analogy to support
St. Hilaire’s hypothesis of transverse deduplication, the similar orientation
of the vascular bundles in the petal and the stamen before it must, as
Celakowsky insists, be good evidence that these represent independent
leaves, and not superposed portions of one.

The main objection to the second hypothesis (that of a suppressed
circle outside of the antipetalous stamens) is that this missing circle,
whether of petals or stamens, is not actually met with in any nearly re-
lated forms (for in Monsonia the fifteen stamens are otherwise explained) ;
also that there are transitions, as above mentioned, between obdiplostemony
and direct diplostemony. To Braun’s theory that the glands behind the
antisepalous stamens in true Geraniacee answer to suppressed phylla,
Eichler objects that these are present behind all ten stamens in Oxalidee ;
also that all are wanting when the oflice of nectar-secretion, which they sub-
serve, is undertaken by some other part of the flower, as by the calyx-spur
in Pelargonium and Tropeolum. The first objection is forcible: the second
mixes morphological considerations with functional, and is inconclusive.
Abortive organs, preserved for their utility as nectaries, might totally dis-
appear when rendered useless by a different provision for the same function.

INCREASED NUMBER OF PARTS. 201

is even more true of greatly multiplied stamens and pistils, as
in Magnolia and Liriodendron, most Anonacez, Ranunculus,
Anemone, and the like. But in Aguilegia, where the number five
is fixed in the perianth, the cyclic arrangement with alternation
of whorls prevails throughout

369. The definite angmentation of calyx and corolla by the
production of one additional whorl of each, and the seeming
anteposition which comes of it when the andrecium remains
simply diplostemonous (in the manner of the Berberidacee,
Menispermaceze, &¢c., 359) has already been explained.

370. Similar increase to two whorls affecting the corolla only
characterizes Anonacezr, Magnoliaceze, Papaveraceze, and Fuma-
riacee. In all but the last order, this is accompanied by indefi-
nitely multiplied stamens, and mostly by an increased number of
earpels. In Fumariacez, which has dimerous flowers, there is a
diminution by the suppression in most cases of half the normal
andreecium, and also an augmentation of the other half by
chorisis. (372.)

371. Parapetalous Multiplication. Under this head may be
described an anomalous arrangement of augmented stamens
which prevails in the order Rosacez, 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 Cratzegus coccinea and
Crus-galli. The ten are in one circle and in pairs, the pairs
alternate with the petals. Some would say the pairs are before
the petals ; but the space between two stamens before each petal
is mostly rather wider than in the pair taken the other way.
The next case in order, as in 15-stamened Hawthorns, and
constantly in Nuttallia, adds to the above a simple interior circle
of five stamens, one directly before the middle of each petal.
Next, as in most Pomez and many Potentillez, there are twenty
stamens, thus placed, but with an additional circle of five alter-
nating with the preceding one. Next there are 25 in three
circles, the second circle as well as the first having ten stamens ;
and finally there are from 30 to 50, all probably in circles of ten
each. ‘There is little doubt that the circles develop in centri-
petal order; the inner successively the later.”

1 It was first clearly described by Dr. A. Dickson, in Trans. Bot. Soc.
Edinb. viii. 468, and Seemann’s Jour. Bot. iv. 473 (1866). He introduced the
term, parapetalous, which is characteristic of it in its elementary form (254,
note): it is particularly illustrated by Eichler, in Bliithendiagramme, ii.
495-510. ‘The former interprets it by chorisis, both median and collateral :
the latter presents the facts and possible views, but declines to adopt either
of them.

2 Accordingly, the whole is probably to be explained by some modifica-

202 THE FLOWER.

372. Chorisis or Deduplication. Both these terms, and the
ideas which they denote, originated with Dunal, but were first
expounded by Moquin-Tandon.’ The first word is Greek for a
separating or separation. The second is a translation of Dunal’s
French word dédoublement (literally undoubling), the ambiguity
of which, and of the original presentation of the case, long
retarded the right apprehension of the subject. Diremption has
been suggested (by St. Hilaire) as a proper term. The mean-
ing simply is, the division of that which is morphologically
one organ into two or more (a division which is of course
congenital), so that two or more organs occupy the position of
one. As thus used, chorisis is restricted, or nearly so, to the
homologues of leaves in the flower, and mainly to stamens and
carpels ; the division or splitting up of a petal or a sepal, when it
occurs, being expressed in the phrases which are applied to leaves.
Yet a compound leaf, especially one of the palmate type, is
a good type of chorisis, the several blades of a compound leaf
answering to the single blade of a simple leaf. It has been ob-
jected against the terms chorisis and deduplication that they
assume the division of that which has never been united; but
so equally does the established terminology of foliage. A di-
vided leaf has never been entire.

373. Chorisis is complete when the parts concerned are dis-
tinct or separate to the very insertion, as in the stamen-clusters of
Hypericum. The foliar form of this would be represented by

—

tion of the augmentation of circles. Dickson’s hypothesis, that the two,
three, or five stamens which are more or less in face of each petal are all
deduplications of that petal, would come to be noticed under the next head,
but it may be dismissed at once. Yet that the pairs in the outer circle
represent each an antisepalous stamen, divided by chorisis (sometimes
incompletely) and much separated, is not improbable. The other tenable
explanation (which may be harmonized with the last) is that the outer
circle of stamens here rightly consists of ten members, respectively alternat-
ing with the sepals and petals taken asa whole. This makes them para-
petalous, and at the same time brings them under Hofmeister’s general law
that new organs originate over intervals of those preceding, in this case over
the ten perianth-intervals directly. It also accords with Hartog’s elucidation
of the accessory parts in the flower of Sapotacez (in Trimen’s Jour. Bot. 1878).
The inner circles are there sometimes 5-merous after the primitive type,
sometimes 10-merous in regular alternation to the preceding circles.

1 Moquin-Tandon, Essai des Dédoublemens, &c., Montpellier, 1826; Con-
sidérations sur les Irrégularités de la Corolle, &c., in Ann. Sci. Nat. xxvii.
237, 1832; Tératologie Végétale, 337. Dumnal, Essai sur les Vacciniées,
1819, cited by Moquin (some pages printed, but never published) ; Considé-
rations sur la Nature et les Rapports de quelques-uns des Organes de la
Fleur, 1829. The next botanist to develop it was St. Hilaire, Morphologie
Végétale, 1841.

CHORISIS OR DEDUPLICATION. 203

such sessile palmately compound leaves as those of some species
of Aspalathus. It is incomplete when division does not extend
to the base; as in Fig. 387, 395. Compare, as a
proximate homologue of this, a petal of Mignonette,
Fig. 385. But proper chorisis requires that the
supernumerary organs should be developed like
unto the original organ which is thus multiplied, or
should complete their symmetry, whatever it be.

374. St. Hilaire distinguished two kinds of deduplication; viz.,
collateral when the members stand side by side, and parallel
when an organ becomes double or multiple antero-posteriorly.
The latter, sometimes called vertical, and sometimes transverse,
is better named median chorisis. The collateral is the origi-
nal and typical chorisis. Most botanists incline to restrict the
name to this, and to give some other explanation and name to
the median form of augmentation. But some cases, such as
those of Tilia and Sparmannia, are clearly of the same nature
as the collateral, and may be a disguised form of it; there are
others which may be explained in accordance with it; and there
are such transitions between some of these and coronal out-
growths that the term chorisis is most conveniently made to
comprise augmentation or doubling in either plane. Distinct
anteposition, however, may be explained in other ways. (357.)

375. Typical or Collateral Chorisis, in which the members,
together answering to one leaf, normally stand side by side,
occurs in many families of plants, and
in a variety of forms. <A few are here
presented.

376. Elodes Virginica (a common
marsh plant of the Hypericum family),
like most of its near relatives, has its
calyx and corolla on the plan of five,
its stamens and carpels on the plan of at pe
three, as is shown in the diagram, Fig. 386. This makes a break
in the symmetry between the corolla and the stamens; but all
within is in regular alternation when the three stamens of each
cluster are counted as one as their union at base into a phulanr
(Fig. 387) may suggest. These phalanges alternate with the
three carpels, and therefore stand where single stamens belong.
The three conspicuous green projections, which in a general way

FIG. 385. A petal of Mignonette (Reseda odorata), with many parted blade,
enlarged. -

FIG. 386. Diagram of flower of Elodes Virginica, with three phalanges of stamens
forming the inner circle, and three glands answering to the outer circle. 387. A de-
tached phalanx of three stamens.

204 THE FLOWER.

are called glands, alternate with the phalanges, and so are taken
to represent the outer circle of stamens. The morphologist
accordingly sees in the glands the homologues or representatives
of the outer series of stamens, reduced to three by abortion, and
in the three stamen-clusters only the three alternating stamens
of the inner series, trebled by chorisis, and this chorisis incom-
plete, because it has not quite divided the filament into three.
In Hypericum, the glands are completely suppressed, each pha-
lanx is almost or quite divided into a cluster, either of about
three stamens each, as in H. Sarothra, or of a few more (in H.
mutilum and H. Canadense), or of an indefinite number, as in
the common St. Johnsworts. Then in some other species (as
in our H. pyramidatum) the carpels and the stamen-clusters rise
to five, realizing complete pentamerous symmetry, except that
the almost numberless stamens all belong to the one inner circle.
Morphologically, they are comparable to the leaflets of five (or
in most species three)
decompound and _ ses-
sile or almost sessile
leaves. The indefinitely
numerous stamens of
Ricinus are similarly
increased from five by
compound ramification.

377. Fumariacex, the
Fumitory family, may
furnish the next illus-
tration. The flower is
on the plan of two
(dimerous) throughout.

ZS

S-
oe

392 393
Taking Dicentra to show it, there is first a pair of small and
scale-shaped sepals, not unlike the pair of bractlets on the

FIG. 388. Dicentra Cucullaria (Dutchman’s Breeches), ascape in flower and a leaf,
severed from the singular bulb (formed of the enlarged bases of petioles). 389. Detached
flower, of natural size, showing also the pair of bractlets on the pedicel. 390. Same
with parts displayed, and 391, inner petals placed above. 392. Diagram of flower of
Dicentra or Adlumia, from a section across the summit. 393. One of the phalanges of
stamens of Adlumia; upper part only.

CHORISIS OR DEDUPLICATION. 2O5

pedicel below (Fig. 389, 390), and normally alternate with them :
alternate with these is a pair of large petals, deeply saccate or
spurred below; alternate with these, a pair of smaller petals
with spoon-shaped tips which cohere at the apex (the corolla
therefore of two circles asin the related Poppy family) ; alternate
with these, two phalanges or united stamen-clusters, of three
stamens each; alternate with these is nothing, for the second
set of stamens is wanting; alternate with this vacancy is a pair
of carpels wholly combined into a compound 2-merous pistil.
The statement itself explains the morphology. ‘The three sta-
mens of each phalanx stand in the place of a stamen, and are
the divisions of one. In Dicentra the members of the phalanx
are almost separate; in Adlumia (Fig. 393) and Corydalis the
undivided filament reaches almost up to the anthers. The middle
anther of the phalanx is normal, or two-celled; the lateral
anthers are one-celled, as if halved.?

1 Eichler adopts this interpretation (proposed in Gray, Gen. Illustr.
i. 118), and applies it to the crucial instance of Hypecoum. In the flower
of this Old World genus, there are four apparently simple and complete
stamens, one before each petal: the simplest interpretation would be that
which the facts appear to present, viz. that both dimerous circles of stamens
are complete and normal. But Eichler —in view of the early development
and the double vascular bundles of the stamens before the inner petals, and
some occasional slight disjunction of their anther-cells — considers that
the interior stamen-circle is wanting here, no less than in the other genera
of the order; that what here takes its place before each inner petal is a
stamen composed of the adjacent lateral member of the phalanx, congeni-
tally severed from the group to which it belongs and soldered into one fila-
ment, bearing the two one-celled anthers so brought together as to imitate a
normal two-celled anther. The organogeny of the blossom is thought to
favor this hypothesis; and it certainly favors the view here adopted of
the composition of the three-membered phalanx of the family generally.
If this interpretation of Hypecoum seems far-fetched, it is no more so than
its exact counterpart, through which DeCandolle, Lindley, and others explain
the case of the rest of the family. Starting with that genus as the simple
type, they conceive that the stamen opposed to each inner petal is each
severed into two, and that these half-stamens attached to the sides of the two
intact stamens, thus producing the phalanges by coalescence.

A good empirical conception of the formation, from a single leaf, of three
stamens in Fumariacez, or two in Crucifere, is afforded by the petals of
Hypecoum, as illustrated by Eichler. The outer petals are slightly three-
lobed from the apex; the inner are deeply so and narrower. The mem-
bers of the next circle in the family generally are just such three-lobed
bodies, the tip of each lobe transformed into an anther. There is an ap-
parent congruity in the production by the symmetrical middle lobe of a
symmetrical two-celled anther, and of a one-celled anther by each unsym-
metrical lateral lobe or stipule-like portion. A fuller development of these
sides of the leaf, and non-development of the middle portion (somewhat
after the analogy of Lathyrus Aphaca, Fig. 219), with anther-formation,
would convert the leaf into a pair of stamens.

206 THE FLOWER.

378. The obvious relationship of Cruciferee to Fumariacee,
their agreement in the rare peculiarity of having the two carpels
595 side by side instead of fore and aft
(median), and the characteristic
anomaly which the andreecium pre-
sents (7. e. the tetradynamy), would
give reason to expect that its prob-
lems might be solved by chorisis.
Indeed, the doctrine was applied to
this, long before its application to
the other order. Beginning at the
centre (Fig. 395, &c.), the pistil is
of two carpels, right and left ; alter-
nate with these is a pair of stamens
on the side next the axis, matched
by another pair on the opposite
side of the pistil, the four longer
and interior stamens; alternate
with these, and lower in insertion,
397 a single stamen on each side ; next,
four petals, of somewhat various overlapping in eestivation,
which essentially alternate with the two single stamens and the
two pairs; lastly, four sepals, alternating with the four petals as
a whole, the anterior and posterior overlapping the lateral ones
in the bud. Now the median (7. e. the anterior and posterior)
pairs of stamens occasionally have their contiguous filaments
conjoined, as in Fig. 397. If this were at all constant, the
inference would undoubtedly be that the case is one of chorisis,
and that the flower as to its essential organs is dimerous. ‘This
is apparently the best explanation to be given. It assumes that
the chorisis is normally complete in the andreecium of Cruciferee,
instead of incomplete, as in Fumariaceze.t And this view is
confirmed by the fact that the median stamens are simple and

1 The hypothesis here adopted, as to the andrecium, is that of Steinheil
(1839), and of Eichler (in Flora, 1865, 1872, and Bliithend. ii. 200), replacing
that of Kunth, 1833, &c., employed in former editions. The rejected view
makes the flower 4-merous up to the pistil, and the stamens all of one circle.
alternating with the four petals, the median stamens (as in our view) doubled
by chorisis. Krause and Wretschko (cited as above by Eichler) would
have the floral circles 2-merous and 4-merous by turns; the calyx of two
2-merous circles (which it plainly is); the corolla of one 4-merous circle

FIG. 394. A cruciferous flower. 395. Diagram of such a flower, with position of
axis marked aboveit. 396. Tetradynamous stamens and the pistil. 397. A common
monstrosity of the same, two of the four inner stamens combined into a common
2-antheriferous body.

CHORISIS OR DEDUPLICATION. 207

single in Senebiera and many species of Lepidium, in which the
lateral or short stamens are at the same time abortive.

379. It is quite possible that chorisis may be extended to the
corolla of the cruciferous flower, and reduce the whole to a
symmetrical 2-merous plan, and to congruity in the perianth
also with Fumariaceze. The only obstacle is in the petals form-
ing a whorl of four where all the rest is 2-merous, for the sepals
are manifestly two decussating pairs. Now the median petals
of Hypecoum are deeply 3-lobed. An abortion of their middle
lobe would leave them almost two-parted: a little more would
separate them ; then they would imitate the four cruciferous petals
as in the diagram, Fig. 395. Applying this view to Crucifere,
the blossom in the two orders would accord in having a 2-merous
three-whorled perianth, the first and third whorls median ;? as
also in the dimerous androecium, the first whorl of which is
lateral. ‘The difference is that in Fumariacez the two members
of the first whorl of stamens augment by chorisis into three, and
the second is wanting, or is present only in Hypecoum ; while in
Cruciferze the first whorl is simple (of the two short stamens), and
the second is doubled. In Fumariacez only the first whorl of the
perianth counts as calyx, and
the corolla is of two whorls ;
in Cruciferze, the first and
second whorls are calyx, the
inner sepals answering to the
outer petals of Fumariacee.

380. Chorisis along with
anteposition of stamens is well
seen in Tilia or Linden, at least in the American species. In
these the indefinitely numerous stamens are in five clusters, one
before each petal (Fig. 398, 399), and there is a petal-like body

399

alternating with the calyx-members as a whole; the short stamens following
as a 2-merous circle; then the long stamens as a 4merous circle; lastly the
2-merous gynecium. G. Henslow (in Trans. Linn. Soc. ser. 2, i. 195) would
have the flower 4-merous by the suppression of the fifth members of a
5-merous type, and a further suppression of half of the remaining exterior
stamen-circle, &c. Finally, there is the much better-maintained view that
the cruciferous flower is 2-merous throughout, as explained in the following
paragraph, 379.

1 This view was taken by Steinheil, in Ann. Sci. Nat. ser. 2, 337 (1839),
and is essentially reproduced by a Russian botanist, Meschajeff, in Bull.
Soc. Imp. Nat. Mosc. 1872.

FIG. 398. Diagram of the flower of Tilia Americana, the common American Lin-
den or Basswood.
FIG. 399. A detached stamen-cluster with its petal-like scale.

298 THE FLOWER.

in each cluster with which the stamens cohere. The explanation
by chorisis is that each cluster, petal-like body included, is a
multiplication of one stamen. The diagram (Fig. 398) accu-
rately shows that most of the stamens originate from the outer
side of the base of the petal-like portion: this is most naturally
explained by median chorisis. The superposition of the clusters
to the petals will take the same explanation as that of Rhamnus,
Vitis, &c. (Fig. 363.) That the andreecium is here composed
of the inner circle merely is partly confirmed by the alternation
of the carpels with the clusters. According to Duchartre,’ the
development of the andreecium in a Mallow indicates a similar
structure; for the whole united mass originates from five protu-
berances, one before each forming petal and connected with it,
this by collateral chorisis forming a cluster of stamens, and the
five clusters coalescing as they develop into a tube of filaments,
such as in Fig. 485. Now Hibiscus and its near relatives have
a naked tip to the stamen-tube, ending usually in five teeth ;
and Sidaleea, as is most strikingly shown in the Californian
S. diploscypha, has two series of stamens, the outer (answering
to those of Malva and its relatives) in five membranaceous pha-
langes, superposed to the petals; the rather numerous inner
series, more or less in phalanges, surmounts an interior filament-
tube. Whence it is inferred
that these, and the five teeth
terminating the column in
Hibiscus, represent the in-
ner stamineal circle which is
wanting in Malva, as it is in
Tilia.?

381. The case of Parnas-
sia would be explained as
analogous to that of Tilia,
but with the stamen-clusters before the petals wholly sterile,
and of fewer divisions, while an inner circle of five stamens

1 Comptes Rendus, 1844, & Ann. Sci. Nat. ser. 3, iv. 123. Duchartre and

others who draw freely upon median chorisis to explain anteposition, and
consider that congenital union proves it, take the phalanges in these cases,
like the single stamens in Vitis, to be an inner part of the petal itself. But
this view appears to have had its day.

2 Gray, Gen. Illustr. ii. 44, 57, 75-82. The position of the carpels before
the petals in Pavonia and Malvaviscus brings the former into symmetrical
alternation with such an inner stamen-circle; but it is not so in Hibiscus,
which has the carpels before the sepals.

FIG. 400. A petal of Parnassia Caroliniana, with a triple staminodium before it.
FIG. 4091, Diagram of the flower of Parnassia Caroliniana.

OUTGROWTHS. 209

alternate with the petals forms the effective andreecium. For
the scale-like body before each petal, and even slightly adnate
to its base (in P. Caroliniana about 3-parted, as in Fig. 400, but
in P. palustris a thin scale, fringed with more numerous gland-
tipped filaments), is plainly outside the stamens in the full-grown
flower-bud. But Eichler and Drude have found
that it is inside in the early bud.* Wherefore, if
these stamen-like bodies really represent a circle
of the andrecium, it must be the inner one; and
that is the more probable view.

382. Multiplication by chorisis in the gyncecium
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
Malvacez, the same thing takes place in Pavonia
and its allies: while in Malope and two other
genera of the same order the few normal carpels are multiplied,
evidently by chorisis, into an indefinite number of wholly distinct
ones.

§ 8. OUTGROWTHs.

383. Proper chorisis is the congenital multiplication of one
organ into two or more of the same nature and office; or at
least into two or more organs, even if dissimilar, as in the
American Lindens, in which one member of the cluster is a kind
of petal. Between this and the production by an organ of ap-
pendages, or outgrowths of little or no morphological signifi-
cation, there are many gradations; as also between these and
mere cellular outgrowths from the surface, even down _ to
bristles and hairs. The latter, in all their variety and modifica-
tions, are properly outgrowths of the epidermis only, and there-
fore consist of extended cells, single or combined, unaccompanied
by vascular or woody tissue. To them has been given the
general name of Zrichomes (Trichoma, pl. trichomata), that is
structures of which hairs are the type. They may occur upon
the surface of any organ whatever. Their morphology is the
morphology of cells rather than of organs. They will therefore
be most conveniently illustrated under Vegetable Anatomy as

1 Wichler in FI. Brasil., Sauvagesiace, & Bliithend. ii. 424; Drude in Lin-
nea, xxxix. 239. Eichler refers to this as a confirmation of Celakowsky’s
explanation of obdiplostemony by posterior displacement. (365.)

FIG. 402. Pistil of Drosera filiformis with tricarpellary ovary (transversely divided),
and six styles, i. ¢. three, and each two-parted.

210 THE FLOWER.

respects their structure, and in the Glossary as respects ter-
minology.

384. But into some bristles, such as those of Drosera, a sub-
jacent stratum of tissue enters, including one or more ducts or
even some woody tissue. Prickles are of this class; and from
the most slender, which pass into bristles, there are all grada-
tions of stoutness and induration. Such outgrowths may even be
formed in most regular order, as the prickles on the calyx-tube
of Agrimonia and scales on the acorn-cup of Oaks, and yet have
no morphological importance. On the other hand, true represen-
tatives of leaf or stem may, by abortion and depauperation, be
reduced to the structure as well as the appearance of trichomes.
Examples of this are familiar in the pappus (answering to limb
of the calyx) of many Compositz, and in the bristles which
answer to perianth in many Cyperacez. The scarious stipules
of Paronychia and of Potamogeton, the ligule of Grasses, and
even the corolla in Plantago, are equally reduced to mere cellular
tissue. So that the structural difference between trichomes and
ouigrowths* is not at all absolute, and the morphological distine-
tion must rest upon other ground than anatomical structure.

385. Among the corolline outgrowths most akin to chorisis is
the Crown (Corona) of Silene and allied Caryophyllacez, at the
junction of the claw
with the blade of
the petals (Fig. 403),
the analogy and
probable homology
of which to the ligule
of Grasses (Fig.
150) is evident ; also
the many-rayed fila-
mentous crown of
Passion-flowers (Fig. 404), which consists of two or more series
of such outgrowths. In Sapindus and some other Sapindaceze,
these ligular outgrowths or internal appendages are more like
a doubling of the petal; as also in Erythroxylum, where they

1 This is the best English name for the Emergenzen of the Germans, the
Epiblastema of Warming, &c. For the development and discussion of this
subject, see Warming, in Kjébenhavn Vidensk. Meddel. 1872, and a larger
treatise on Ramification in Phanerogams, Copenhagen, 1872. Also, Uhl-
worm in Bot. Zeit. 1873; Celakowsky in Flora, 1874; and Eichler’s note on
Emergenzen in Bliithendiagramme, i. 48.

FIG. 403. Petal of Silene Pennsylvanica, with its crown.
FIG. 404. Flower of Passiflora ce@rulea, reduced in size.

FORMS OF THE RECEPTACLE. 211

are often more complicated in structure. They are always on
the inner face, and are commonly two-lobed or parted.

386. Similar stamineal appendages are well known in Cuscuta
(Dodder), in Larrea (Fig. 405) and other Zygophyl-
laceze, and less conspicuously in Gaura.

387. To extend to them the name of LictLte may
not be amiss, whether they are regarded as mere
outgrowths of floral leaves, without further morpho-
logical relations, or whether they be, at least some-
times, interpreted as the homologue of intrapetiolar
stipules, as their ordinarily two-cleft form, and their
coincidence in Erythroxylum with an intrapetiolar two-cleft stipule
suggest.

§ 9. Forms or THE Torus OR RECEPTACLE.

388. Torus is the more specific and proper name, RECEPTACLE
is the more usual. (303.) A normal receptacle of the flower
would be that of Fig. 316, the apex of the flower-stalk somewhat
enlarged, roundish or depressed, and with surface mainly cov-
ered by the insertion of the several organs; the Be inter-
nodes which it potentially contains being
undeveloped. As the members of the flower
multiply and occupy numerous ranks, the
receptacle enlarges or lengthens to give them
insertion or standing-room.

389. Of elongated forms of receptacle,
Magnolia and Liriodendron or Tulip-tree give
familiar instances. The lengthening in the
former is mainly for the support of both an-
dreecium and gyncecium ; in the latter, as in
Myosurus, mainly for the gyncecium only.
The fall of the matured carpels reveals it
as a very slender or bodkin-shaped pro-
longed axis. Of broadened forms, the Straw-
berry, even in blossom, affords a familiar
example. (Fig. 406.) In the same order,
Rubus odoratus shows a very broad and flat
receptacle: in roses, it is so deeply concave as to become the
reverse of the strawberry (Fig. 407), being urn-shaped with a
narrow mouth, upon which the petals and stamens are borne,

FIG. 405. Stamen of Larrea Mexicana, with a couspicuous ligulate appendage at
the base within.

FIG. 406. Receptacle of a strawberry in longitudinal section. 407. Same ofa rose,
in diagram.

912 THE FLOWER.

while the pistils line the walls of the cavity, the base or centre
of this cavity answering to the apex of the strawberry.

390. Sometimes internodes are lengthened between certain
members. In Schizandra, the receptacle, barely oblong in blos-
som, lengthens greatly in fruiting, so as to scatter the carpels
on a long filiform axis.

391. In many Gentians, in Stanleya and Warea among
Cruciferze, and in most species of Cleome, the internode of the
receptacle between stamens
and pistil is developed into
a long stalk to the latter.
Gynandropsis (Fig. 409) is
like its near relative, Cleome,
except that this very long stalk
has the lower part of the
stamens adnate to it: the in-
ternode between the corolla
and calyx is broad and slightly
elevated (or in Cleome, &e.,
narrower and longer) ; and

He iF so the several floral circles
are as it were spaced apart by this unusual development of
receptacular internodes. In Silene (Fig. 408) and many other
plants of the Pink family, an internode between the calyx and
corolla is prolonged into a stalk or Stipe."

1 Srrpe is the general name of a stalk formed by the receptacle or some
part of it, or by a carpel. To distinguish its particular nature in any case,
the following terms are more or less employed : —

THECAPHORE, for a stipe which belongs to a simple pistil itself (where
it is homologous with a petiole), and is no part of the receptacle, as in Coptis
or Goldthread.

GyNopruHore, where the stipe is an internode of receptacle next below the
gynecium, as the pod-stalk in some Cruciferae, Cleome, and Gynandropsis.

GonopPuoreE, when it elevates both stamens and pistil, as it seemingly
does in the lower stipe of Gynandropsis, Fig. 409.

ANTHOPHORE, when the stipe is a developed internode between the
calyx and corolla, as in the Pink family, Fig. 408.

GYNOBASE is a term properly applied to a short and comparatively broad
portion of receptacle on which the gynecium rests, as in Rue and Orange
(Fig. 414), Houndstongue, Sage, &c. This may extend up between the car-
pels and pass into, or the upper part become a

CARPOPHORE, a name properly applied to a portion of receptacle which
is prolonged between the carpels as a central axis, as in Geranium (Fig.
411) and many Umbellifere, Fig. 412.

FIG. 408. Section of a flower of Silene Pennsylvanica, showing the stipe or
anthophore.

FIG. 409, Flower of Gynandropsis, with floral circles separated on the elongated
receptacle.

FORMS OF THE RECEPTACLE. 213

392. Instead of forming a stalk, the elongation may be continued
between the carpels in the form of a slender axis, as in Gera-
nium (Fig. 410, 411), and in the carpophore
of the fruit of Umbelliferee, Fig. 412. In
Geranium, this prolongation of receptacle

410

4i4

extends far above the ovaries as a beak, to which the styles are
adnate for most of their length.

393. In Nelumbium (Fig. 415), the gynophore, or portion of
receptacle above the stamens, is enlarged into a singular broadly
top-shaped body, with a flat summit, in which the pistils (a dozen
or more isolated carpels) are separately immersed.

394. A Disk is a part of the receptacle, or a development of it,
enlarged under or around the pistil. When under it or around
its base and free from the calyx,
the disk is hypogynous, as in
Orange, Fig. 414. Here it is a
kind of gynobase. When adher-
ent to or lining the base of the
calyx, it is perigynous, as in Mae
Rhamnus (Fig. 415,416) and Cherry (Fig. 357): when carried
by complete adnation up to the summit of the ovary, it is ep7gy-
nous, as in Cornus, in Umbelliferee, &c. Not rarely it divides
into lobes, as in Vitis (Fig. 379, 380), in Periwinkle and most
Apocynaceous plants, and in Cruciferee. These are termed g/ands
of the disk, and indeed are commonly glandular or nectariferous.

FIG. 410. Gyneecium of Geranium maculatum. 411. Thesame with fruit mature,
the five ovaries or cells and the lower part of their styles separated and recurving away
from the prolongation of the axis or receptacle, to which they were at flowering-time
firmly attached.

FIG. 412. Mature fruit of Osmorrhiza. the two carpels splitting away below from
the filiform prolongation of the receptacle, or carpophore.

FIG. 413. The top-shaped receptacle of Nelumbium, with the pistils, immersed in
hollows of its upper face.

FIG. 415. Flower of a Rhamnus or Buckthorn, and 416, section of the same, show-
ing a thickened perigynous disk.

214 THE FLOWER.

It is not possible by any direct demonstration to distinguish be-
tween such productions of the receptacle, which are classed as
belonging to the axis, and suppressed or undeveloped phyllous
organs, such as stamens, which glands of the disk may some-
times represent.

395. Hypanthium. Inspection of Fig. 415, 416, and 337, and
comparison with Fig. 339, will suggest an explanation differ-
ent from that which is generally
adopted. Instead of regarding the
calyx as beginning on a level with
the base of the ovary, and the cup
as lined, more or less thickly, by
an expansion of the receptacle (the
perigynous disk),
the calyx may be
understood to begin
where this and the
ovary become free
from each other.
Underthat view, the
receptacle, instead
of convex or protu-
berant, is here con-
cave, has grown up
around the ovary,
which, however, is free from the cup in the earlier cited figures,
but immersed in it in Fig. 339 and the like. A comparison with
a rose-hip, an apple, and a pear much strengthens this interpre-
tation, which is rather largely adopted at this day,-at least
theoretically. It was perhaps first proposed by Link, who intro-
duced the appropriate name of Hypanrnum. A hypanthium or
hypanthial receptacle is, as the name betokens, a flower-axis or
receptacle developed mainly under the calyx. The name is a
good one, in any case; and such structures as those of Calycan-
thus (Fig. 417-419), a rose, a pear (the lower part of which is
evidently an enlargement of peduncle), and of Cactus-flowers
(Fig. 317), although quite compatible with the theory of adnation,
are more simply explained by it.?

1 But, whether the cases are well distinguishable or not, it by no means
follows that the receptacle plays such a part in all instances of perigyny and
of inferior or partly inferior ovary. Such a view is attended by more diffi-
culties than the other. Unless the mediation of an invisible receptacle must

FIG. 417. Flowering branch of Calycanthus. 418. Vertical section of the urn-shaped

receptacle, the imbricated bracts or sepals on its surface cut away. 419. Mature
fructiferous receptacle entire, showing some scars from which the bracts have fallen.

ADAPTATIONS TO FERTILIZATION. 215

Section IV. Certrars ADAPTATIONS OF THE FLOWER TO THE
ACT OF FERTILIZATION.

§1. In GENERAL.

396. The introduction into morphological botany of the con-
siderations now to be mentioned should have dated from the
year 1793, in which Christian Conrad Sprengel published his
curious treatise on the structure of flowers in special reference
to insect aid in their fertilization. For this book, which was
wholly neglected and overlooked for more than sixty years, con-
tains along with some fanciful ideas the germs of the present
doctrine and many excellent illustrations of it.!. The interest in
the doctrine now prevalent is witnessed by a copious special
literature, beginning with the publication, in 1862, of Darwin’s
book on the fertilization of Orchids by the aid of insects.?

be invoked whenever there is a junction of two dissimilar organs, the petals
and stamens of a Lythrum or a Cuphea are united with the calyx itself,
instead of calyx beginning at the top of a long and simple tube. And if
three or more of the floral whorls may be congenitally united, why not these
also with the remaining one? Van Tieghem, in his Anatomie Comparée de
la Fleur, maintains wholly the old view, founding it upon anatomical struc-
ture and his ability to trace down to the base of the ovary the distinct
vascular bundles of the several involved organs.

1 C. C. Sprengel, Das entdeckte Geheimniss der Natur im Bau und der
Befruchtung der Blumen, Berlin, 1793. Even earlier, Kelreuter ( Vorlaufige
Nachricht, ete., 1761-1766) recognized the necessity of insect-aid to various
blossoms, and described some special contrivances for the purpose.

2 Charles Darwin, On the Various Contrivances by which British and
Foreign Orchids are fertilized by Insects, and on the Good Effects of Inter-
crossing, London, 1862. Ed. 2, 1877. This last contains a list of the papers
and books which bear upon the subject, published since 1862.

Other leading works and papers on the subject are, exclusive of the
other volumes and papers of Darwin, more or less referred to hereafter.

Treviranus, Ueber Dichogamie, &c., in Bot. Zeitung, xxi. 1863.

Hugo gon Mohl, Einige Beobachtungen iiber dimorphe Bliithen, Bot.
Zeitung, xxi. 1863.

Delpino, Pensieri sulla Biologia Vegetale, &c., 1867. Relazione sull’
Apparecchio della Fecondazione nelle Asclepiadie, &c., 1867. Ulteriore
Osservazioni sulla Dichogamia, &c., 1868-69, 1870, and later papers.

Axell, Om anordningarna for de Fanerogama Vaxternas Befruchtung,
Stockholm, 1869.

Hildebrand, Die Geschlechter-Vertheilung bei den Pflanzen, 1867, and
other papers.

Hermann 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 Professor Kerner, which describes arrangements in blossoms for exclud-
ing unwelcome guests, has not yet reached us. It introduces the new terms
Autogamy and Allogamy, defined on the following page; the latter compre-

216 THE FLOWER.

397. The subject, here considered as a part of morphology,
must be fully treated, as regards acts and processes, under physi-
ology. Every thing in the flower is in relation to fertilization
and fructification, directly or indirectly. This section is con-
cerned with those adaptations of structure by means of which
agents external to the blossom are brought into service for its
fertilization.

398. Linnzeus and his successors taught that the adjustments
in hermaphrodite flowers were such, on the whole, as to secure the
application of the pollen of its stamens to the stigma of its pistil
or pistils. The present view is, that this is doubtless strictly
secured in certain flowers of a moderate number of species, but
never in all the flowers of any such species; that in ordinary
flowers, where it may commonly take place, it is not universal ;
that in the larger number of species there is something or other
in the floral structure which impedes or prevents it. Some
flowers are adapted for close fertilization ; some for cross fertili-
zation; some for either. Here two terms need definition, viz. :

Close fertilization or Self-fertilization, or Autogamy, the applica-
tion and action of a flower’s pollen upon its own pistil ;

Cross fertilization, or Allogamy, the action of the pollen of one
flower on the pistil of some other flower of the same species.
This may be near, as when between flowers borne in the same
cluster or on the same plant; remote, when between flowers of
distinct plants of the same immediate parentage ; most remote,
when between different races of the same species. Any thing
beyond this is hybridization, or crossing of species.

§ 2. ADAPTATIONS FOR ALLOGAMY OR INTERCROSSING.

899. The doctrine now maintained appears to have been first
propounded by Sprengel in the statement that ‘* Nature seems
to have wished that no flower should be fertilized by* its own
pollen,” —a proposition which is not wholly tenable, for there
are blossoms specially adapted to self-fertilization. It was re-
affirmed in our day by Darwin, in a similar adage, ‘* Nature
abhors perpetual self-fertilization,’”” — a metaphorical expression
to which no effective exception has been taken. And the infer-
ence was drawn by him, that some important good to the species
must result from propagation through the union of distinct
individuals, and especially of individuals which have been dis-
tinct for several or many generations.

hending Geitonogamy, fertilization by pollen of other flowers of the same
plant, and Xenogamy, by pollen from a flower on another plant.

ADAPTATIONS FOR INTERCROSSING. yr} i

400. The actual proposition, simply stated, is that flowers are
habitually intercrossed, and that there are manifold structural
adaptations which secure or favor intercrossing, to such extent
as to justify the proposition. The 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
array of special adaptations for which no other sufficient reason
appears, or (to resume the metaphor) from the vast pains which
seem to have been taken to secure this end. ‘This inference has
been to some extent confirmed by direct experiment.*

401. Separation of the sexes is a direct adaptation to inter-
crossing, rendering it necessary between individuals in dicecious,
and largely fay oring it in most moncecious and polygamous
flowers. Strictly close fertilization can occur in hermaphrodite
flowers only ; but it is in these that the most curious adaptations
for intercrossing are revealed.

402. The agencies to the one or the other of which most
flowers are structurally adapted in reference to intercrossing are
mainly two; viz., the winds and animals, of these chiefly insects.
Delpino has accordingly classified flowers into Anemophilous and
Entomophilous ; literally wind-lovers and insect-lovers, but de-
noting wind-fertilized and insect-fertilized, according to the
agent by which pollen is transported.* There are hermaphrodite
and unisexual flowers of both classes, but most wind- ee
flowers are unisexual.

403. Wind-fertilizable or anemophilous flowers are mostly neu-
tral or dull in color, destitute of odor, and not nectariferous.
Their principal structural adaptations to this end, besides the
separation of the sexes in most of them, are the superabundance,
incoherency, dryness, and lightness of the pollen, rendering it
very transportable by wind and currents of air. The immense
abundance of pollen, its lightness, and its free and far diffusion
through the air in Pines, Firs, Taxodium, and other Conifers,
are familiar. Their pollen fills the air of a forest during anthe-
sis; and the ‘* showers of sulphur,” popularly so-called, the
yellow powder which after a transient shower accumulates as
a scum on the surface of water several or many miles from the

1 Darwin, The Effects of Cross and Self-Fertilization in the Vegetable
Kingdom, London, 1876. American Edition, New York, 1877.

2 Ornithophilous, i.e. bird-fertilized, flowers are to be ranked with entomo-
philous. The large blossoms of Trumpet Creeper (Tecoma radicans) and
of Trumpet Honeysuckle (Lonicera sempervirens), and others, are commonly
visited and probably fertilized by humming-birds as well as by moths; and
other birds are known to play a similar part in equatorial regions.

218 THE FLOWER.

nearest source, testifies to these particulars. All amentaceous
trees (Willows excepted), Hemp, Hops, &c., are wind-fertilized ;
and, among perfect flowers, those of most Grasses, Sedges, and
Plantago. In the latter families especially, the anthers are pro-
truded or hung out in the air only when just ready to discharge
their pollen, and are at that moment suspended on suddenly
lengthened capillary drooping filaments, fluttering in the gentlest
breeze ; and the stigmas are either dissected into plumes, as in
most Grasses, or beset with copious hairs on which pollen is
caught. One physiological adaptation, very common in the fol-
lowing class, is not unknown among hermaphrodite wind-fertiliz-
able flowers, where it is important for securing intercrossing, viz.
Dichogamy. It is best seen in the common species of Plantago
or Plantain, and is described below. (408.)

404. Insect-fertilizable or entomophilous flowers are correlated
with showy coloration (including white, which is most showy at
dusk), odor, or secretion of nectar, often by all three modes of
attraction to insects combined. Some insects, moreover, visit
flowers for their pollen, a highly nutritious article, and ordina-
rily produced in such abundance that much may be spared.
The showiness of corolla or other floral envelopes is an attractive
adaptation to fertilization, enabling blossoms to be discerned at
a distance; nor do we know-that fragrance or other scent or
that nectar subserves any other uses to the flower than that of
alluring insects. Adaptations in the pollen of such blossoms
for transportation by insects are various. Commonly the grains
are slightly moist or glutinous, or roughish, or studded with
projections, or strung with threads (as in CSnothera), so as not
to be readily dispersed in the air, but to have some slight
coherence as well as capability of adhering to the head, limbs,
or bodies of insects, especially to their rough surfaces; and
in two families (Orchidaceze, Asclepiadacez) the pollen is com-
bined in masses and with special adaptations for being trans-
ported en masse. (421.) With this the stigma is usually
correlated, by roughness, moisture, or glutinosity.!

405. Adaptations of the flower itself in reference to insect
visitation are wonderfully various ; and most of these are found
upon investigation to favor, or often to necessitate, intercross-
ing. In dicecious flowers, this is necessitated by the separation ;
in moneecious and polygamous flowers, of various kinds and

1 Thus nearly every Orchid genus but one has a persistently glutinous.
stigma ; in the exceptional one, Cypripedium, it is moist and minutely rough-
ened, in correlation with the loosely granular or pultaceous pollen which it
is to receive.

ADAPTATIONS FOR INTERCROSSING. 219

degrees of separation, pollen is very commonly borne from
plant to plant; in hermaphrodite flowers only are more special
arrangements needed to secure intercrossing ora certain measure
of it, and in these such arrangements abound.

406. Irregularity is one of the commonest modifications of the
flower (326, 337): it is never conspicuous except in blossoms
visited by insects and generally fertilized by their aid; and it
finds rational explanation on the score of utility in this regard.*

407. Dichogamy, a term introduced by C. C. Sprengel, who
first noticed and described it, is one of the most usual and effect-
ual (rather physiological than morphological) adaptations for the
promotion of intercrossing between hermaphrodite flowers. It
means that such intercrossing is brought to pass by a difference
in the time of maturity of anthers and stigma; this rendering
dichogamous blossoms practically the same as dicecious or mon-
cecious in respect to fertilization, while there is the economical
gain that all the flowers are fertile. According to whether the
anthers or the stigmas are precocious, dichogamous flowers are

Proterandrous (or Protandrous), when the anthers mature and
discharge their pollen before the stigma of that blossom is recep-
tive of pollen ;

Proterogynous (or Protogynous), when the stigmas are in
receptive condition before the anthers have matured their pollen.

Synanthesis,? the maturing of the two sexes simultaneously or
nearly so, is however made to secure the same result through
special arrangements.

408. Proterogyny. The Plantains, such as Plantago major and
P. lanceolata, are familiar instances of this in a wind-fertilized
genus with hermaphrodite flowers. The anthesis proceeds from
base to apex of the spike in regular order, and rather slowly.
While the anthers are still in the unopened corolla and on short
filaments, the long and slender hairy stigma projects from the tip
and is receiving pollen blown to it from neighboring plants or

1 This did not escape the attention of Sprengel in the last century, and
slong with it the fact that strictly terminal and also vertical flowers, whether
erect or suspended, are seldom irregular, while comparatively horizontal or
obliquely set flowers more commonly are so. The irregularity is in refer-
ence to a landing place for the visiting insect, or also to storage of or accessi-
bility to nectar, &e.

Darwin (Forms of Flowers, 147) remarks that he does not know of a
single instance of an irregular flower which is wind-fertilized.

2 Synacmy is the term proposed by A. W. Bennett, in Journal of Botany,
viii. (1870), 316, with its opposite, Heteracmy, for proterandry and proterogyny.
The latter names, in their shorter form (protandry and protogyny), appear to
have originated with Hildebrand, 1867.

220 THE FLOWER.

spikes: a day or two afterwards, the corolla opens, the filaments
greatly lengthen, and the four anthers now pendent from them
give their light pollen to the wind ; but the stigmas of that flower
and of all below it on that spike are withered or past receiving
pollen. Among Grasses, Anthoxanthum is in the same case.
The arrangement is somewhat similar to the Plantain in Amor-
pha, which is fertilized by insects, the simple stigma projecting
beyond the corolla in bud, while the anthers are still immature

420

and enclosed. Scrophularia is a good instance of proterogyny
in flowers fertilized by bees. The flower is irregular (Fig.
420422), and is approached from the front, the spreading lower
lobe being the landing place. Fig. 420 represents a freshly
opened blossom ; and Fig. 421, a section of it. Only the style
tipped with the stigma is in view, leaning over the landing place ;
the still closed anthers are ensconced below. The next day or
a little later all is as in Fig. 422. The style, now flabby, has
fallen upon the front lobe, its stigma dry and no longer receptive :
the now-opening anthers are brought upward and forward to the
position which the stigma occupied before. A honey-bee, taking
nectar from the bottom of the corolla, will be dusted with pollen
from the later flower, and on passing to one in the earlier state
will deposit some of it on its fresh stigma. Self-fertilization
here can hardly ever take place, and only through some disturb-
ance of the natural course.

409. Proterandry. The process is the reverse, and is at-
tended with much more extended movements in Clerodendron
Thompsoniz, a Verbenaceous tropical African climber now com-
mon in conservatories. The adaptations in this flower (which
we indicated long ago) are exquisite. The crimson corolla
and bright white calyx in combination are very conspicuous.
The long filiform filaments and style, upwardly enrolled in the

FIG. 420, 421. Early opened flower of Scrophularia nodosa, and a longitudinal
section. 422. Flower a day or two later.

ADAPTATIONS FOR INTERCROSSING. aA |

bud, straighten and project when the corolla opens: the stamens
remain straight, but the style proceeds to curve downward and
backward, as in Fig. 423. The anthers are now discharging
pollen: the stigmas are immature and closed. Fig. 424 repre-
sents the flower on the second day, the anthers effete, and the
filaments recurved and rolled up spirally; while the style has

taken the position of the filaments, and the two stigmas now
separated and receptive are in the very position of the anthers the
previous day. The entrance by which the proboscis of a butterfly
may reach the nectar at bottom is at the upper side of the orifice.
The flower cannot self-fertilize. A good-sized insect flying from
blossom to blossom, and plant to plant, must transport pollen
from the one to the stigma of the other.

410. Proterandry abounds among common flowers. It is
conspicuous in Gentians and in nearly all that family. But.
while in Gentians the short style is immovable and erect, in
Sabbatia it is thrown strongly to one side, out of the way of and
far below the stamens, the branches closed and often twisted, so
that the stigma is quite inaccessible until the stamens have shed
their pollen: then the style becomes erect, untwists, its two flat
branches separate, and expose the stigmatic surface of their inner
face in the place which the anthers occupied. In Sabbatia
angularis, Lester F. Ward’ observed that the anthers of freshly

1 In Meehan’s Gardeners’ Monthly, September, 1878, 278.

FIG, 423. Flower of Clerodendron Thompsoniz, first day; 424, second day.

222 THE FLOWER.

opened blossoms are all thrown to one side almost as strongly
as the style is thrown in the opposite direction. One of our
common Fireweeds, Epilobium angustifolium or E. spicatum, as
it is variously called, which is common all round the northern
hemisphere, is similar to Sabbatia in behavior. In the freshly
opened flower, while the anthers are in good condition and are

giving their pollen to bees, the still immature style is strongly
curved downward and backward, as in Fig. 425. Two or three
days later, when the pollen is mostly shed, the style straightens,
lengthens to its full dimensions, and spreads its four stigmas over
the line of the axis of the blossom (Fig. 426), in the very
position to be pollinated by a bee coming from an earlier flower.
411. In the following instances of proterandry, the style is
made the instrument of distributing the pollen which it is not
itself to use. The
j anthers of a Cam-
panula discharge all
their pollen in the
unopened bud, and it
is nearly all deposited
on the style which
they surround, the
upper part of which
is clothed with a coat
of hairs for holding
= ies the pollen. (Fig. 427.)

In the open flower, the stamens are found to be empty and withered,
as in Fig. 428. These flowers are visited by bees and other insects
for the pollen. While this is going on, and while the pollen is
fresh and plentiful, no stigma is apparent. Later, the top of the
style opens into three (in some species five) short and spreading
branches, the inner faces of which are the stigmas. Although

FIG. 425, 426. Flowers of Epilobium angustifolium or spicatum; in the first,
freshly expanded; in the second, a few days older.

FIG. 427. Vertical section of an unopened flower of Campanula rapunculoides:
the broad white lines are sections of two anthers. 428. Same of an older flower.

ADAPTATIONS FOR INTERCROSSING. 223

so close at hand, little if any of the pollen of that flower can
reach the stigmas. These actually get fertilized by pollen
brought by bees, which come loaded with it from :
other flowers and other plants. Symphyandra differs
from a true Campanula chiefly in the continued
cohesion of the five anthers into a tube around the
style. (Fig. 429, 430.) The pollen is discharged on
and held by the hairy upper portion of the style.
Soon after, the corolla expands, the lower
part of the style lengthens, and carries
the pollen-loaded part out of and above
the anther-tube, as in Fig. 430; lastly, the
three connivent, tips of the style diverge
and expose the stigmas to pollen mainly
brought by bees from other flowers. By a
slight further modification in Lobelia and
in Composit, pollen is pushed out of the
anther-tube by the tip of the style as it
lengthens, or by the very back of the two
stigmas, the faces of which, afterwards
exposed, are not to receive this, but other
pollen, though it may at times receive some of its own. The
arrangement in Composite is here illustrated from Leptosyne
maritima (Fig. 431-435), a showy plant of Southern California,
now not very rare in cultivation. The large flowers around the

FIG.-429. Stamens and pistil of a young, and 430, same from an old flower of
Symphyandra pendula.
FIG. 431. Head of flowers of Leptosyne maritima, of the natural size.

pct 8 THE FLOWER.

margin (ray-flowers, with ligulate corolla), one of which is sepa-
rately shown in Fig. 432, are pistillate only: the enlarged and
extended open part of the
corolla (bright yellow in
color) serves for attrac-
tion, the circle of rays
gives the appearance as
of a single large flower.
The flowers of the disk
or whole central part are
hermaphrodite, and with
narrow tubular corollas,
from the orifice of which
projects the greater part

“ of the tube of five co-
alescent anthers. The pollen is early discharged into the interior
of this tube. The style, with somewhat enlarged and brush-like
tip, at first reaches only to the
bottom of the anther-tube: it
slowly lengthens, pushes the
pollen before it out of the tube
(Fig. 433) and into the way of
insects of various kind, which,
travelling over the surface, con-
vey it to older flowers of the same

head and of other plants. The style, elongating yet more, raises
some of the pollen still higher (as in Fig. 434) ; and at length its
two branches separate and diverge (Fig. 435), exposing to other
pollen the stigmatic receptive surface which until now was un-
approachable.

412. In Parnassia, which has sessile stigmas, their receptive
surface is actually not formed until the anthers become effete ;

FIG. 432. A ligulate female flower of the same, and a central hermaphrodite flower.
433. Upper part of the latter, more enlarged, the tube of anthers projecting from the
corolla, and the pollen projecting from apex of the anther-tube, being pushed up by the
lengthening of the style beneath. 434. This style now projecting, and some pollen still
resting on its tip. 435. Tip of same style (more advanced and magnified); the two
branches spreading, still carrying some pollen on the apex of each arm or branch, and
by the “ivergence now exposing the stigmatic inner faces.

ADAPTATIONS FOR INTERCROSSING. 225

and, as the plants or stems are single-flowered, they are function-
ally dicecious while structurally hermaphrodite.

413. The adaptations for hermaphrodite intercrossing with
synanthesis (407), 7. e. where there is no essential difference of
time in the maturing of anthers and stigma, are manifold.
They may be classed into those without and those with dimor-
phism of stamens and pistils, or, in other words, those with
Homogonous and those with Heterogonous flowers.}

414. The cases without dimorphism are the most various,
certain families having special types; and are of all degrees,
from those that require intercrossing to those that merely favor
or permit it. For the present purpose, having only morphology
in view, it suffices to bring to view two or three cases or types of

415. Particular Adaptations in hermaphrodite blossoms, not
involving either dichogamy or dimorphism. These are exceed-
ingly various; but they may be distinguished into two general
kinds, namely: 1, where loose and
powdery pollen is transported from
blossom to blossom in separate grains,
and 2, where pollen-masses or the
whole contents of anthers are bodily
so transported.

416. Papilionaceous flowers (such
as pea-blossoms, 358) — 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. ser. 5, xiii. 82, and in Amer.
Naturalist, January, 1877. Dimorphism in flowers may affect the perianth
only, and not the yovj or essential organs; or there may be two kinds of
flowers as respects these also, but with no reciprocal relations, as in cle/sto-
gamous dimorphism (534) ; or of two kinds essentially alike except in stamens
and pistil, and these reciprocally adapted to each other, which is heterogonous
dimorphism, or, when of three kinds, trimorphism.

FIG. 436. Flower of Wistaria Sinensis natural size. 437. Same enlarged, with
standard, wings, and half the keel removed. 438. Same with the keel depressed, as it
is when a bee alights on this its usual landing place, the cluster of authers and stigma
thus brought up against the bee’s abdomen. 439. Style and stigma, with part of the
ovary, more magnified, a fringe of fine bristles around the stigma.

15

226 THE FLOWER.

Wistaria (Fig. 456-439), in which the light fringe of stiff hairs
around the stigma (shown in Fig. 439) would not prevent pollen
of surrounding anthers from falling upon it. Yet when a bee
alights upon the keel,
with head toward the
base of the flower, and
proboscis is inserted
for nectar between the
foot of the standard
and the keel, the latter
is depressed by the
weight, so that the ab-
' domen of the insect is
brought against the ten
anthers and the stig-
ma, becoming thereby
smeared with pollen,
some of which when
other blossoms are vis-
ited cannot fail to be applied to their stigmas. ‘The very similar
flower of Locust (Robinia), like that of the Pea, adds an adapta-
tion in favor of intercrossing. The style for some length below the
stigma is covered with a short beard of hairs, as is
seen in Fig. 442. The anthers open early and dis-
charge their pollen, which mainly lodges on this
beard (Fig. 443), in a manner which may thus far
be likened to the case of Campanula. (411.) The
wings and the keel are yoked together, and are
together depressed by the weight of an alighting

443 bee. This does not bring out the anthers as in
Wistaria, but these remain untileffete within the sac, while the
stigma and the pollen-laden part of the style (Fig. 441) are
projected against the bee’s abdomen, which, by the oblique
movement, is first touched by the stigma and next brushed over
with pollen by the style below. So that, in visiting a succession
of blossoms, some pollen of one flower is transferred to the body
of the bee, and thence to the stigma of the next flower, which
flower immediately gives to the same spot some of its pollen, to
be transferred to the next flower’s stigma, and so on.

417. Two special modifications of the papilionaceous type

FIG. 440. Flower of Robinia hispida, the standard and wings removed. 441. Same,
as depressed by the weight of a bee, causing the stigma and pollen-laden tip of the style
to protrude. 442. Enlarged section of same in the bud, leaving one keel-petal, half the
stamens, and the pistil in view. 443. Style and stigma at a later period, the beard
loaded with pollen; more magnified,

ADAPTATIONS FOR INTERCROSSING. 227

need particular mention. One of them, the Bean-blossom, is
weil known to botanists; the other not so. The peculiarity in
the common Bean, Phaseolus vulgaris, and its nearest relatives,
is that the keel, enclosing the stamens and pistil, is prolonged

into a narrow snout which is spirally coiled (as in Fig. 444-446) ;
that the stigma is oblique on the tip of the style, and the beard
on the style is mainly on the same side that the stigma is: the
wing-petals stand forward and turn downward, forming a con-
venient landing place for bees. As in the Locust-blossom, the
anthers early discharge their pollen, much of which adheres

lightly to the beard of the style. In the untouched flower, all
from first to last is concealed in the coiled keel. Press down the
wing-petals, and first the stigma and then the pollen-laden tip
of the style projects from the orifice : remove the pressure, and
they withdraw within. When this pressure is made by a bee,
resting on the wing-petals while searching for nectar within the
base of the blossom between the keel and the standard, the same
movement occurs: the stigma first, and then the pollen on the
style, strikes against a certain portion of the front or side of
the bee’s body, and the repetition of this operation causes the
fertilization of each blossom by other than its own pollen. A
slighter pressure or lighter movement of the wing-petals suffices

FIG. 444. Flower of Garden Bean, Phaseolus vulgaris. 445 Same with wing-
petals pressed down and tip of style projecting from the orifice of the keel. 446. Same
as 444 enlarged, and standard and wings remoyed. 447. Upper part of keel, in the
condition of 445, enlarged, showing plainly the projecting style. 448 Section of the
keel, enlarged, showing the style within before the anthers open: stamens for sake of
clearness not delineated. 449. Pistil detached from an older flower; the brush loaded
with pollen.

228 THE FLOWER.

to jostle some of the pollen down upon its own stigma, so that
self-fertilization is not uncommon.

418. Apios tuberosa, a near relative of Phaseolus, exhibits a
different and equally curious modification of the same parts.
The wing-petals for landing place
are similar: the standard is pro-
portionally large, firm in texture,
and shell-shaped or concave, with
a small boss at the tip as seen from
behind, or a shallow sae as seen

ees from the front: the keel is narrow
and sickle-shaped ; it arches across the front of the flower, and
the blunt apex rests in the notch or shallow sac of the tip of the

452 453
standard. (Fig. 450, 452,453.) So it remains if untouched until
the blossom withers: no self-fertilization has ever been observed,
and none ordinarily occurs. The anthers
are assembled close around the stigma,
but a little short of it (Fig. 452); the
pollen is not early nor copiously shed in
the enclosure: the small terminal stigma
is at first covered with a pulpy secretion,
which at length collects into a soft ring
around its base over or through which no
pollen passes. But when the Keel is liber-
ated by lifting from underneath, it curves
promptly into the shape shown in Fig.

FIG 450. Flower of Apios tuberosa, unvisited. 451. Same after visitation, the
keel dislodged from the retaining notch, and more incurved; the tip of the style pro-
truded and thrust forward, followed by the anthers.

FIG. 452. Enlarged vertical section of flower-bud of Apios tuberosa. 453 A flower
with half the standard cut away, to show the blunt apex of the keel resting in the
notch. 454. Diagram of flower, with half of the standard cut away, toshow what takes
place when the apex of the keel is liberated. The figures (also those from 423 to the
present), and the first account of the adaptations of Apios, were published in the Amer-
ican Agriculturist in 1876.

ADAPTATIONS FOR INTERCROSSING. 229

451, or better in Fig. 454, where the dotted lines indicate its
original position; and first the end of the style, tipped with
its stigma, is pushed forward, and then the anthers come into
view. The flowers are visited by humble-bees, and sometimes
by honey-bees. In searching for nectar at the base of the flower,
they probably push forward into the space under the arching
keel, and by slightly elevating dislodge its apex ; when first the
stigma and then the anthers are brought against some portion
of the insect’s body, and against the same portion in succeeding
blossoms, thus effecting cross-fertilization. This rationally ex-
plains a remarkable adaptation, which seems to be not otherwise
intelligible.

419. Special “Adaptations. Two of these, each peculiar to
the genus, may here be referred to. In Kalmia-blossoms (Fig.

457

=
455-458), the anthers discharge the pollen through a small
orifice at the apex of each cell, in this respect agreeing with
Rhododendrons and their other relatives; but
none of them utilize this family peculiarity in the
manner of Kalmia. In the flower-bud, each of the
ten anthers is lodged in a small cavity or pocket
(externally a boss) of the corolla, in a way analo-
gous to that in which the keel of Apios is lodged in
the tip of the standard (418) : the expansion of the
border of the corolla in anthesis curves the fila-
ments outward and backward ; and when the bowed — 4s

stamens are liberated by rough jostling they fly up elastically,
and the pollen is projected from the two orifices. Some pollen
may possibly be thrown upon the single small stigma at the
tip of the style, which rises much above the stamens. But the
anthers are not dislodged when undisturbed, at least until
after the elasticity of the filaments is lost: they are dislodged by
humble-bees, which circle on the wing over the blossom, the

FIG. 455. Vertical section of a flower-bud of Kalmia latifolia, showing the anthers
lodged in the pockets of the corolla. 456. Expanded flower, with bowed stamens
457. Vertical section of the same. 458. A stamen, enlarged.

250 THE FLOWER. :
under side of the abdomen frequently touching the stigma, while
the proboscis is searching round the bottom of the flower, liberat-
ing the stamens in the process, which one by one project their
pollen upon the under side of the insect’s body. In the passage
from flower to flower, pollen is thus conveyed from the anthers
of one to the stigma of another.

420. Iris has three stamens, one before each sepal or outer
lobe of the perianth, and behind each petal-like lobe of the style
(Fig. 459): the stigma, a
shelf-like plate of each lobe,
is just above the anther;
but, as the anther faces
outward and the stigma is
higher and faces inward, no
pollen can find its way from
the one to the other. But the
adaptation of parts is admir-
able for conveyance by bees,
which, standing upon the
only landing place, the re-
curved sepal, thrust the head
down below the anther, and
in raising it carry off pollen,
to be afterwards lodged
upon the stigmas of other
flowers which they visit.

421. Transportation of Pollinia, or of all the pollen in a
mass, is effected in most of the species of two large orders,
not otherwise allied, the Asclepiadaceze and the Orchidacee.
While in the Iris family the number of stamens is reduced from
six to three, in all the Orchis family, except Cypripedium, the
stamens are further reduced to a single one; but the pollen is
peculiarly economized. That of Arethusa is in four loose and
soft pellets, in an inverted casque-shaped case, hinged at the
back, resting on a shelf, the lower face of which is glutinous
stigma, over the front edge of which the casque-shaped anther
slightly projects ; and this anther is raised by the head of a bee
when escaping out of the gorge of the flower. The loose pellets
of pollen are caught upon the bee’s head, to the rough sur-
face of which they are liable to adhere lightly and so to be carried
to the flower of another individual, there left upon its glutinous

FIG. 459. Flower of Iris pumila, with front portion and half of one petaloid style-
lobe and stigma cut away. The section of the stigma is seen edgewise: the rough
upper surface only is stigmatic.

ADAPTATIONS FOR INTERCROSSING. 231

stigma by the same upward movement which immediately after-
ward raises the anther-lid and carries away its pollen, to be
transferred to a third blossom, and so on.

422. But it isin Orchis
and in the commoner re-
presentatives of Orchis in
North America (viz. Ha-
benaria, &c.) that the
most exquisite adapta-
tions are found, and the
greatest economy — se-
cured ; paralleled, how-
ever, by most of the very
humerous and various epi-
phytic and by various ter-
restrial Orchids of warmer
regions. <A single illus-
tration may here suffice ;
and Darwin’s volume on
the Fertilization of Or-
chids (596, note), with
its references to the
copious literature of the
subject, may be studied
for full particulars and
their bearings. The flower
is trimerous, and the peri-
anth adnate to the ovary,
therefore apparently de-
veloped upon its sum-
mit. The three external
parts of the perianth,
which in Habenaria orbi-
culata (Fig. 460) are
much the broader, are the
sepals: the three alternate
and internal, the petals :
the base of the long and narrow petal which is turned downward
is hollowed out and extended below into a long tube, closed at
bottom, open at top (the spur or nectary), in which nectar is

1

1
Y

FIG. 460. Flower of Habenaria or Platanthera orbiculata, enlarged. 461. Combined
stamen and stigma, more enlarged. 462. One of the two pollen-masses (pollinia), with
its stalk and glutinous disk or gland. 462¢. Lower part of this stalk and its disk, more
magnified.

232 THE FLOWER.

copiously secreted and contained. The central part of the
blossom, beyond the orifice of the nectary (shown separately in
Fig. 461), consists of one anther and a stigma, fused together
(the clinandrium): the marginal portions, opening by a long
chink, are the two cells of the anther, approximate at their
broader portion above, widely divergent below: most of the
lower part of the space between is excessively glutinous, and is
the stigma. The grains of pollen are united by means of short
threads of very elastic tissue into small masses, and these into
larger, and at length into pellets, having stalks of the same
B elastic tissue, by which they are

all attached to a firmer central
stalk, or caudicle. (Fig. 465-465.)
To the lower end of this caudicle
(directly to the end of it in our
Habenariz and Orchises gener-
ally, in this instance to the inner
side of the end, with a thick inter-
mediate base intervening), is at-
tached a button-shaped disk, the
face of which is exposed, and is
on a line with the surface of the
anther; so that these two disks
look toward each other across the
broad intervening stigmatic space,
as seen in Fig. 461. The exposed
face of the disk being covered with
463 465 a durable layer of very viscid mat-

ter, the body itself is sometimes termed a gland, and not improperly.
The viscidity is nearly of the same nature as that of the interven-
ing stigma, of which the glands are generally supposed to be
detached portions. If so, then a portion of the stigma is cut off
from the rest and specialized to the purpose of conveyance of the
pollen. When a finger’s end or any smaller body is touched to
these disks, they adhere so firmly that the attached pollinia or
pollen-masses are dragged out of the cell and carried away en-
tire. Some of these pollen-masses have been found attached by
the disk to the eyes of a large moth. When a moth of the size
of head and length of proboscis of Sphynx drupiferarum visits a
spike of these flowers, and presses its head into the centre of the

FIG. 463. A more magnified pollen-mass of Platanthera orbiculata, with its stalk
and gland. 464. Five of the separate portions or pollen-packets, with some of the
elastic threads of tissue connecting them. 465. A portion more highly magnified, with
some of the pollen-grains in fours detached.

ADAPTATIONS FOR INTERCROSSING. 233

flower so that its proboscis may reach and drain the bottom of
the nectariferous tube, a pollen-mass will usually be affixed to
each eye: on withdrawal,
these will stand as in Fig.
466. Within a minute they
will be turned downward
(Fig. 4667), not by their
weight, but by a contraction
in drying of one side of the
thick piece which connects
the disk with the stalk.
When a moth in this con-
dition passes from the last
open flower of one spike to
that of another plant, and
thrusts its proboscis down a
nectary, the transported pol-
len-masses will be brought
in contact with the large
glutinous stigma: on with-
drawal, either some of the
small pellets of pollen will be
left adherent to the stigma,
the connecting elastic threads
giving way; or else a whole
pollen-mass will be so left,
its adhesion to the glutinous aoe
stigma being greater than that of the disk to the moth’s eye.
The former is a common and a more economical proceeding, as
then a succession of flowers are abundantly fertilized by one
or two pollen-masses. In either case, new pollen-masses are
earried off from fresh flowers and applied to the fertilization
of other blossoms on the same and eventually on those of differ-
ent individuals. Cases like this, and hundreds more, all equally
remarkable, serve to show how sedulous. sure. and economical are
the adaptations and processes of Nature for the intercrossing of
hermaphrodite flowers.

422¢, An arrangement analogous to that of Orchids, and
similarly subservient to cross-fertilization, characterizes the
otherwise widely unlike Asclepias family. In Asclepias (Milk-
weed) there are five stamens surrounding a large stigmatic

FIG. 466. Front part of Sphynx drupiferarum, bearing a pollen-mass of Platan-
thera orbiculata affixed to each eye, in the early position. 466. Front view of the head,
later, showing the pollen-masses deflexed.

oa

Zot THE FLOWER.

body, and alternating with these five two-cleft glands, the ver-
tical chink or groove of which is glucinous. To each gland is
firmly attached, by a caudicle or stalk, a pollen-mass of an ad-
jacent anther. (Fig. 522.) <A slight force upraising the gland
detaches it from the stigma and drags the pair of suspended
pollen-masses out of their cells. Insects visiting the blossoms
commonly dislodge them, the gland adhering to their legs or
tongues when these happen to be drawn through the adhesive
chink, and convey them from one flower to another. Without
such aid the flowers of Asclepias ravely set seed.'

423. Dimorphism, 7. e. the case of two kinds of blossoms, both
hermaphrodite, on the same species, is another adaptation to
intercrossing. Not all dimorphism, however, for in eleistogamous
dimorphism (484) the intent to self-fertilize is evident. There
may also be dimorphism as to the perianth, not particularly
affecting fertilization. One kind, however, and the commonest,
is a special adaptation to intercrossing, viz. :

424. Heterogonous Dimorphism. (415, note.) This term is
applied to the case in which a species produces two kinds of
hermaphrodite flowers, occupying different individuals, the flowers
essentially similar except in the andreecium and gyneecium, but
these reciprocally different in length or height, and the adapta-
tions such that, by the agency of insects, the pollen from the
stamens of the one sort reciprocally fertilizes the stigma of the
other.?, This dimorphism has been detected in about forty genera
belonging to fourteen or fifteen natural orders, widely scattered
through the vegetable kingdom ; but there are far more examples
among the Rubiacez than in any other order. Sometimes all
the species of a genus are heterogonous, as in Houstonia, and

1 The reported sensitiveness of the gland, referred to in the first issue of
this volume (1879), was founded upon misinterpreted observations.

2 This peculiar arrangement has been long known in a few plants, such
as Primula veris, P. grandiflora, and Houstonia. In Torrey and Gray’s
Flora of North America, ii. 38, 39 (1843), these flowers are said to be dicecio-
dimorphous, not’ denoting that they are at all unisexual, but that the two
forms occupy different individuals. Their meaning was detected by C.
Darwin, and made known in his paper “On the Two Forms or Dimorphic
Condition in the Species of Primula, and on their. Remarkable Sexual Rela-
tions,” published in the Journal of the Linnean Society, vi. (1862), 77: repub-
lished, in 1877, as the leading chapter of his volume entitled “The Different
Forms of Flowers on Plants of the Same Species.” Mr. Darwin had termed
these flowers simply Dimorphic; but in this volume he adopted Hilde-
brand’s name of Heterostyled for this kind of blossom. The difference,
however, affects the andreecium, and even the pollen, as well as the style;
wherefore we proposed for it the name of Heteroyonous or Hetereyone dimor-
phism, as mentioned in a former note, 413.

ADAPTATIONS FOR INTERCROSSING. 235

Cinchona, sometimes only a part of them, as in Primula and
Linum. In Hottonia, a Primulaceous genus of two species, the
European one has heterogonous dimorphism’ for cross-fertiliza-
tion: the American one has homogonous showy flowers with
only the general chance for intercrossing, and earlier flowers
which are cleistogamous for self-fertilization.

425. The nature of heterogone dimorphism may be well under-
stood from a single example. The most familiar one is that of
Houstonia ; but, in larger blossoms, Gelsemium is a fine illus-
tration in the Southern United States, and Mitchella (Fig. 467)
mostly in the Northern. Raised from the seed, the individuals

are about equally divided between the two forms: namely, one
form with long style and short or low-inserted stamens; the
other with short style and long or high-inserted stamens. The
stigmas in one rise to about the same height as the stamens in
the other, both in the tall or exserted organs and in their low
and included counterparts, as is shown in Fig. 468, answering
to the left hand and Fig. 469 to tle right hand flowers of Fig. 467.
A bee or other insect with proboscis of about the length of the
corolla-tube, visiting the blossoms of Mitchella, will brush the
same part of its body against the high anthers of the long-
stamened and the high stigmas of the long-styled forms; and

> - r-O

1 ©. C. Sprengel, as Darwin mentions, had noticed this, before 1793. He,
“with his usual sagacity, adds that he does not believe the existence of
the two forms to be accidental, though he cannot explain their purpose.”
Darwin, Forms of Flowers, 51.

Some heterogonous Primulas are said to produce homogonous varieties in
cultivation. In Primula, and in other genera, there are species which seem
as if of one sort only, no reciprocal sort being known, as if one form had
become self-fertile and the other had disappeared.

FIG. 467. Partridge Berry, Mitchella repens, in the two forms, viz. long-stamened
and short-styled, and short-stamened and long-style. :

236 THE FLOWER.

the same part of the proboscis against the low anthers of the
short-stamened and the low stigmas of the short-styled form.

426. Moreover, Dar-
{i win has ascertained by
microscopical examina-
tion that the pollen of
the two differs in size or
shape, and by experi-
ment that it is less
active upon its own
stigma than upon the
other; indeed, that in
many cases (as in some
species of Linum) it is
quite inactive or impo-
tent not only upon its
own stigma but upon its
own-form stigma, while
it is prepotent on the other, and this reciprocally of the two
forms.’ Here, then, are flowers structurally hermaphrodite, but
functionally as if dicecious, securing all the advantages of the
latter, along with the economical advantage that both sorts of
individual and every blossom may bear seed. With dicecism
only about half the plants could be fruitful.

427. Heterogonous Trimorphism. A threefold heterogonism
is known in certain species of a few genera; and this complica-
tion may have certain conceivable advantages over dimorphism.
Where seedling dimorphous individuals are few and far between
(those multiplying from root would all be alike), there would
be an even chance that any two near each other were of the
same form and therefore sterile or imperfectly fertile. But if
the organization were of three forms, any two of which inter-
crossed with perfect fertility, the chances (as Darwin remarks)
are two to one that any two plants were of different forms, and
therefore by fertilizing each other completely fruitful.

428. The earliest known instance of three forms as to recip-
rocal relative length of stamens and pistil is that of Lythrum

Ss

QS =

1 Impotence of own pollen, either absolute or relative. occurs no less in
certain flowers which are not dimorphous, as in Corydalis, some species of
Passiflora, &c. On the contrary, many dimorphous flowers are in a certain
degree self-fertile, especially in the long-stamened and short-styled form.
These subjects are physiological, and belong to another volume.

FIG. 468. Long-styled flower of Fig. 467, laid open. 469. Long-stamened flower of
the same laid open. Both equally enlarged.

ADAPTATIONS FOR INTERCROSSING. 237

Salicaria. This was indicated by Vaucher in 1841, more par-
ticularly described by Wirtgen in 1848, but was interpreted by
Darwin, and the more recondite differences brought to notice, in
1864.1 ‘+The three forms may be conveniently called, from the
unequal length of their pistils, the long-styled, mid-styled, and
short-styled. The stamens also are of unequal lengths, and
these may be called the longest, mid-length, and shortest.”
The pollen of the different classes of stamens is of two sorts as
to color, and of three as to size, the largest grains from the
largest stamens. ‘* The pistilin each form differs from that in
either of the other forms,
and in each there are two
sets of stamens, different
in appearance and func-
tion. But one set of
stamens in each form ,,...
corresponds with a set
in one of the other two
forms. Altogether, this
one species includes three
females or female organs,
and three sets of male .vica.
organs, all as distinct
from one another as if
they belonged to different
species; and, if smaller.
functional differences are = *!e4
considered, there are five
distinct sets of males. Two of the three hermaphrodites must
coexist, and pollen must be carried by insects reciprocally
from one to the other, in order that either of the two should be
fully fertile; but, unless all three forms coexist, two sets of
stamens will be wasted, and the organization of the species as
a whole will be incomplete. On the other hand, when all three
hermaphrodites coexist, and pollen is carried from one to the
other, the scheme is perfect: there is no waste of pollen and no

1 Tn an article On the Sexual Relations of the Three Forms of Lythrum
Salicaria, in Jour. Linn. Soe. viii. 169. Also on the Character and Hybrid-
like Nature of the Offspring of the Illegitimate Unions of Dimorphic and
Trimorphic Plants. Ibid. x. 393, 1868. Reproduced and extended in his
volume entitled “ Forms of Flowers,” 1877.

FIG. 470. Diagram of the flowers of the three forms of Lythrum Salicaria, in their
natural position, with the petals and calyx removed on the near side. The dotted
lines with the arrows show the directions in which pollen must be carried to each
stigma to ensure full fertility. (From Darwin.)

a

238 THE FLOWER.

false coadaptation.” The whole arrangement is displayed in
the annexed diagram (Fig. 470), and in the following account
of the operation.! ‘* Ina state of nature, the flowers are inces-
santly visited for their nectar by hive and other bees, various
Diptera, and Lepidoptera. The nectar is secreted all round the
base of the ovarium; but a passage is formed along the upper
and inner side of the flower by the lateral deflection (not repre-
sented in the diagram) of the basal portions of the filaments ;
so that insects invariably alight on the projecting stamens and
pistil and insert their proboscides along the upper and inner
margin of the corolla. We can now see why the ends of the
stamens with their anthers and the end of the pistil with the
stigma are a little upturned, so that they may be brushed by
the lower hairy surfaces of the insects’ bodies. The shortest
stamens, which lie enclosed within the calyx of the long- and
mid-styled forms can be touched only by the proboscis and narrow
chin of a bee: hence they have their ends more upturned, and
they are graduated in length, so as to fall into a narrow file,
sure to be raked by the thin intruding proboscis. The anthers
of the longer stamens stand laterally farther apart and are more
nearly on the same level, for they have to brush against the
whole breadth of the insect’s body. . . Now I have found no
exception to the rule that, when the stamens and pistil are bent,
they bend to that side of the flower which secretes nectar.

When nectar is secreted on all sides, they bend to that side
where the structure of the flower allows the easiest access to it,
as in Lythrum. . . . In each of the three forms, two sets of sta-
mens correspond in length with the pistil in the other two forms.
When bees suck the flowers, the anthers of the longest stamens,
bearing the green pollen, are rubbed against the abdomen and
the inner sides of the hind legs, as is likewise the stigma of the
long-styled form. The anthers of the mid-length stamens and
the stigma of the mid-styled form are rubbed against the under
side of the thorax and between the front pair of legs. And,
lastly, the anthers of the shortest stamens and the stigma of the
short-styled form are rubbed against the proboscis and chin; for
the bees in sucking the flowers insert only the front part of their
heads into the flower. On catching bees, I observed much green
pollen on the inner sides of the hind legs and on the abdomen,
and much yellow pollen on the under side of the thorax. There
was also pollen on the chin, and, it may be presumed, on the
proboscis, but this was difficult to observe. I had, however,

1 All from Darwin, Forms of Flowers, 137-147, &.

ADAPTATIONS FOR INTERCROSSING. 239

independent proof that pollen is carried on the proboscis ; for a
small branch of a protected short-styled plant (which produced
spontaneously only two capsules) was accidentally left during
several days pressing against the net, and bees were seen insert-
ing their proboscides through the meshes, and in consequence
numerous capsules were formed on this one small branch. . .
It must not, however, be supposed that the bees do not get more
or less dusted all over with the several kinds of pollen; for this
could be seen to occur with the green pollen from the longest
stamens. . . . Hence insects, and chiefly bees, act both as
general carriers of polien, and as special carriers of the right
sort.”

429. Finally, a long series of experiments (requiring eighteen
distinct kinds of union) proved that both kinds of pollen are
nearly or quite impotent upon the stigma of the same flower, and
that no ovary is fully fertilizable by other than a ‘ legitimate
union,” z.e. by stamens of the corresponding length; but that
the mid-length pistil is more prolific than either of the others
under illegitimate union of either kind; which might perhaps be
expected, as the pollen proper to it is intermediate in size of
grains between that of the long and that of the shortest stamens.

430. Nessa verticillata, a common Lythraceous plant of the
Atlantic United States, is similarly trimorphous, but has not
yet been particularly investigated. Several South African and
American species of Oxalis are equally trimorphous, and have
been investigated by Darwin and Hildebrand,’ with results
quite as decisive as in Lythrum Salicaria. One genus of
Monocotyledons has trimorphous blossoms, viz. Pontederia, of
which the North American P. cordata is a good illustration.’

431. All known flowers exhibiting reciprocal dimorphism or
trimorphism are entomophilous: no such wind-fertilized species
is known. Few of them are irregular, and none very irregular :
they do not occur, for instance, in Leguminosz, Labiate,

1 Monatsber. Akad. Berlin, 1866; Bot. Zeit. 1871, &ce. According to
Darwin, Fritz Mueller “has seen in Brazil a large field, many acres in extent,
covered with the red blossoms of one form [of an Oxalis] alone, and these
did not produce a single seed. His own land is covered with the short-styled
form of another species, and this is equally sterile; but, when the three
forms were planted near together in his garden, they seeded freely.” Forms
of Flowers, 180.

2 Detected by W. H. Leggett. See Bulletin of Torrey Bot. Club, vi. 62,
170; and for the original discovery in Brazilian species, by Fritz Mueller,
see Darwin’s Forms of Flowers, 183, &c. Pontederia has three lengths of
style and counterpart stamens, as in Lythrum Salicaria, each flower having
two sets of stamens, tliree in each set.

240 _THE FLOWER.

Scrophulariaceze, Orchidacerw, &c. Nature is not prodigal, and
does not endow with needless adaptations flowers which are
otherwise provided for.

§ 3. ADAPTATIONS FOR CLOSE FERTILIZATION.

432. Even where cross-fertilization in bisexual flowers is
obviously arranged for, it is apt to be tempered with more or
less of close-fertilization. The more exquisite the arrangements
for the former are, the more completely is the plant dependent
upon insect visitation. Failure to intercross is a remote and
small evil compared with failure to set seed at all. In order
therefore that the plan of cross-fertilization may not defeat even
its own end, through too absolute dependence on precarious
assistance, some opportunity for self-fertilization will usually be
advantageous. Also there is a long array of insect-visited
flowers, especially polyandrous ones, in which close fertilization
must be much the commoner result, except where the pollen of
another but wholly similar flower has greater potency.

433. Subsidiary self-fertilization is secured in a great variety
of ways. In Gentiana Andrewsii, which is proterandrous, and
usually cross-fertilized by humble-bees entering bodily into the
corolla, an exposed surface of pollen long remains fresh upon
the ring of anthers girding the base of the style: when the stigmas
separate, they remain for some days simply divergent, but they
at length become so revolute that the receptive surface is brought
into contact with the ring of pollen below. The opening and
closing of blossoms by day or night, the growth of style, fila-
ments, or corolla after anthesis commences, or other changes of
position, may secure a certain amount of self-fertilization in a
subsidiary or even in a regular way. Then certain species, such
as Chickweed, which blossom through a long season, close-
fertilize even in the bud in early spring, when insects are scarce,
but are habitually intercrossed by insectsin summer. Somewhat
similarly, according to Hermann Mueller.’ certain species, such
as Euphrasia officinalis and Rhinanthus Crista-galli, habitually
produce two kinds of blossoms, one larger and more showy,
usually affecting sunny localities, and with parts adapted to
intercrossing by insects; the other smaller or inconspicuous, and
with anthers adjusted for giving pollen to the adjacent stigma
without aid. There are gradations between these last arrange-
ments, and the more special and remarkable one of dimorphism
with

1 Befruchtung der Blumen durch Insekten, 204; Nature, viii. 433.

ADAPTATIONS FOR CLOSE FERTILIZATION. 241

434. Cleistogamy. Here the intention and the accomplishment
of self-fertilization are unmistakable. This peculiar dimorphism
consists 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 Linnzeus,! 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 Malpighiaceze
in 1832, and recorded in 18435 that Adolphe Brongniart had well
investigated those of Specularia, and that Weddel had discoy-
ered them in Impatiens Nolitangere. <A full account of the then
known cases was given by Mohl? 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
relationship, though all but five are Dicotyiedons. All but the
Grasses*® and Juncus are entomophilous as to the ordinary
flowers, 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 ordinary flowers in such plants
are sterile, and perhaps they always are so except when cross-
fertilized: in most cases they are habitually infertile or spar-
ingly fertile. Probably they suffice to secure in every few
generations such benefit as a cross may give, while the principal

1 Campanula (now Specularia) perfoliata and Ruellia clandestina, the
latter a cleistogamous state of R. tuberosa. Linneus did not make out the
structure of the flowers, but supposed them to want the stamens.

2 In Bot. Zeitung, xxi. 309.

3 In Bot. Zeitung, xvi. 730.

* Ibid. xxv. 65. The name (denoting “ closed up” union or fertilization) has
been written cleistogenous, which is not so proper. We prefer cleistoyamous to
cleistogamic (and so of similar terms), as best harmonizing with the Latin
adjective form, both in form of termination and in euphoniously taking the
accent upon the antepenult.

® Amphicarpum (Milium amphicarpon, Pursh) is the earliest recognized
cleistogamous Grass, except perhaps Leersia oryzoides. Some species of
Sporobolus 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. Sci. January, 1878, 71.

16

»

242 THE FLOWER.

increase is by cleistogamous self-fertilization, which thus offsets
the incidental disadvantage of the former mode.

437. In general, the cleistogamous are like unto the ordinary
flowers arrested in development, some arrested in the almost
fully formed bud, most at an earlier stage, and in the best
marked cases with considerable adaptive modification. In
these, ‘‘ their petals are rudimentary or quite aborted; their
stamens are often reduced in number, with anthers of very small
size, containing few pollen-grains, which have remarkably thin
transparent coats, and generally emit their tubes while still
enclosed within the anther-cells; and, lastly, the pistil is much
reduced in size, with the stigma in some cases hardly at all
developed. These flowers do not secrete nectar or emit any
odor: from their small size, as well as from the corolla being
rudimentary, they are singularly inconspicuous. Consequently,
insects do not visit them; nor, if they did, could they find an
entrance. Such flowers are therefore invariably self-fertilized ;
yet they produce an abundance of seed. In several cases, the
young capsules bury themselves beneath the ground, and the
seeds are there matured. These flowers are developed before,
or after, or simultaneously with the perfect ones.”! In Grasses,
however, as in some Dicotyledons, there is much less modifica-
tion and more transition. For when Leersia half protrudes its
panicle, in the usual way, the included half is fertile and the
expanded portion sterile (or almost always so), although the
flowers may open and exhibit well-developed anthers, ovaries,
and stigmas. But when similar panicles remain enclosed in the
leaf-sheaths, they are mostly fruitful throughout.

438. Fully to apprehend the economy of cleistogamy in pollen-
saving alone,—and contrariwise to estimate the expense of
intererossing, — one should compare the small number of pollen-
grains which so completely serve the purpose in a typical cleis-
togamous flower (say 400 in Oxalis Acetosella, 250 in Impatiens,
100 in some Violets) with the several thousands of all entomo-
philous cross-fertilized flowers, rising to over three and a half
millions in the flower of a Peony, also their still greater number
in many anemophilous blossoms. To this loss should be added
the cost of a corolla and its action, also of the production of
odorous material and of nectar. No species is altogether cleis-
togamous. Thus cleistogamy, with all its special advantage,
testifies to the value of intercrossing.

1 Darwin, Forms of Flowers, 310.

PERIANTH, OR FLOWER-LEAVES. 243

Section V. THe PERIANTH,! oR CAaLyx AND COROLLA IN
PARTICULAR.

439. The distribution of the floral leaves around the axis,
which belongs to phyllotaxy, and their particular disposition in
the bud (zstivation), have already been considered in Chap. IV.
Sect. I., Il. And most of the morphology of calyx and corolla
has been outlined in the preceding sections of the present chap-
ter. What remains chiefly relates to particulars of form and
to terminology.

440. Duration. The differences in this respect give rise to a
few terms, such as the following. Calyx or corolla may be

Persistent, not cast off after anthesis, but remaining unwithered
until the fruit is formed or matured; as the calyx in Labiatee, in
Physalis, and most Roses.

Marcescent, withering or drying without falling away; as the
corolla of Heaths, Drosera, &c.

Deciduous, falling after anthesis and before fructification ; as
the petals of Roses, the calyx and corolla of Columbine.

Ephemeral or Fugacious, lasting for only a day; as the petals
of Poppy, Helianthemum, Purslane, and Spiderwort. In the
two former, they are cast or early deciduous, the anthesis lasting
but a day: in the two latter, the anthesis is equally or more
brief, but the petals deliquesce or decay at once without falling,
as does the whole flower of Cereus grandiflorus and other night-
blooming Cactacee.

Caducous, falling when the blossom opens; as the calyx of
Poppy and Baneberry.

441. Numerical Terms, succinctly denoting the number of
leaves, either of the perianth as a whole, or of any one of its
circles, are common in descriptive botany. The most general
are those which simply specify the number of component leaves,
by prefixing Greek numerals to the Greek name of leaves, ex-
pressing them in Latin form, or transferring them to the Eng-
lish. Thus

Diphyllous, of two leaves (sepals or petals) ; Zriphyllous, of
three ; Tetraphyllous, of four ; Pentaphyllous, of five ; Hexaphyllous,
of six, and so on. A tulip and a Tradescantia flower have a
hexaphyllous perianth, but composed of two circles, answering
to calyx and corolla; each Triphyllous.? When the character

1 Perianthium, alias Perigone or Perigonium. (296.)

2 As elsewhere explained, when numerical composition is indicated without
reference to nature of parts, the terms d/merous, trimerous, tetramerous, penta-
merous, &c., may be used. (522.)

244 THE FLOWER.
of the organ, 7. e. whether calyx or corolla is to be specified,
the word sepal or petal is employed in the combination ; as,

Disepalous, of two sepals ; T'risepalous, of three; Tetrasepalous,
of four; Pentasepalous, of five (also written 5-sepalous, and ac-
cordingly 2-sepalous, 3-sepalous), and so on: also,

Dipetalous, Tripetalous, Tvtrapetalous, Pentapetalous (2-5-
petalous), &c., when the corolla is concerned.

442, Monophyllous, Monosepalous, and Monopetalous are the
proper terms for perianth (calyx, corolla, &c.) composed of a
single leaf. Likewise Polyphyllous, Polysepalous, and Polypetal-
ous for the case of a considerable but unspecified number of
members. Unfortunately, in the Linnean and long-prevalent
use, monopetalous was the term employed to designate a corolla
of one piece in the sense, or the fact, of a coalescence or grow-
ing together of two, three, five, or more petals into a cup or
tube ; and so of a calyx, of a whorl of bracts, &e. And poly-
petalous, polysepalous, and polyphyllous were the counterparts
of this, meaning of more than one distinct piece, whatever the
number. The misleading use, consecrated by long prescription,
is not yet abandoned, but will in time be obsolete. In present
descriptive botany, a polyphyllous calyx, or a_ polypetalous
corolla, or a 5-petalous corolla, would be taken to mean that the
sepals or petals (as the case may be) were distinct or uncom-
bined, and a monopetalous corolla to be one with petals combined
by coalescence. (329.)

443. Terms of Union or Separation. The proper term for a
corolla or a calyx the leaves of which are more or less coalescent
into a cup or tube is

Gamopetalous for such a corolla, Gamosepalous for the calyx ;
these terms meaning united petals or sepals. The older and mis-
leading names JMonopetalous and Monosepalous, although current
up to a recent day, should be discontinued. Another term is
not rarely used in Germany, that of Sympetalous, for the gamo-
petalous (or formerly monopetalous) corolla, — therefore Syn-
sepalous for a similar calyx. It is perhaps a more apt term
than gamopetalous, and of the same etymological signification ;
but the latter is already well in use.

Choripetalous is, on the whole, the most fitting name for a
corolla the petals of which are separate (as it literally expresses
this), that is, for what is still commonly called Po/ypetalous, as
already explained. (442.) Itis adopted by Eichler, &e. Chori-
sepalous is the term applied to the calyx. Déalypetalous (em-
ployed by Endlicher) has the same meaning. Both this term
and choripetalous carry the implication of separated, rather

PERIANTH, OR FLOWER-LEAVES. 245

than of typically separate, parts. Hleutheropetalous (literally
free-petalled) has also been used, but is inconveniently long.

444. Degree of coalescence is most correctly expressed by the
phrases united (connate, or coherent, or coalescent) at the base,
to the middle, or to the summit, 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 in describ-
ing the lobing or toothing of leaves and all plane organs.
(184-188.) That is, the calyx or corolla when gamophyllous
is for description taken as a whole, and is said to be parted
(3-parted, 5-parted, &c.), when the sinuses extend almost to
the base ; cleft, when about to the middle ; /obed, a general term
for any considerable separation beyond toothing ; dentate or
toothed (3-toothed, 5-toothed, &c.), when the union extends
almost to the summit; entive, when the union is complete to
the summit or border.

445. Parts of Petals, &c. The expanded portion of a petal,
like that of a leaf, is the Lamrya or BLape: any much contracted
base is the Uncuts or Ctaw. The latter is very short in a rose-

471 472 473

petal, but long and conspicuous in a pink and all flowers of that
tribe (Fig. 471), in many Capparidez (Fig. 409) and Crucifere.
A sepal is very rarely distinguishable into lamina and claw.
446. Parts of Gamophyllous Perianth. The coalescent portion
of a corolla, calyx, or of a perianth composed of both (such as
a Lily or Crocus-blossom), so far as the sides are parallel or not
too spreading, is its TuBr: an expanded terminal portion, either
divided or undivided, is the Lime or Borper. The limb may

FIG. 471. Corolla of Soapwort, of five separate long-clawed or unguiculate petals,
with a crown at the junction of claw and blade.

FIG. 472. Flower of Gilia coronopifolia; the parts answering to the claws of the
petals of the last figure here all united into a tube.

FIG. 473. Flower of the Cypress-Vine (Ipomca Quamoclit); the petals a little farther
united into a five-lobed spreading border.

FIG. 474. Flower of the Ipomcea coccinea; the five component petals perfectly united
into a trumpet-shaped tube, and beyond into an almost entire spreading border.

246 THE FLOWER.

be parted (that is, the component parts not united) quite or
nearly down to the tube or base, as in Fig. 472, 475 ; or less so,
as in Fig. 473, 476 (with limb
d-lobed) ; or with merely angles
or points to represent the tips
of the component members, as in
Fig. 474; or with even and entire
border, as in common Morning-
Glory, Fig. 482.

447. The line, or sometimes a manifest or conspicuous portion,
between the limb and tube (in the corolla always a portion above
or at the insertion of the stamens, when these are borne by the
corolla) is called the Turoat, in Latin Faux, pl. fauces. This
is mostly more open than the tube, yet less expanded than the
limb ; but it often presents insensible gradations from the one to
the other.

448. Such appendages as the Corona or Crown (385, shown
in Fig. 403, 404, 471) usually belong to the throat of a gamo-
petalous corolla or perianth, as in Oleander, Comfrey, Borrage,
Narcissus, &¢., or to a corresponding position when the parts
are not coalescent.

449. Forms of Corolla, Calyx, &e.- As to terminology, some
of these are special and are applicable to corolla only, as the

Papilionaceous, the peculiar irregular corolla of the typical
portion of Leguminose (388, Fig. 342-334), which has been
already illustrated, and in which the petals, two pairs and an
odd one, take particular names. Also the

Caryophyllaccous, or Fink-flower (Fig. 471), a regular corolla,
of five long-clawed (wnguiculate) petals, the claws enclosed in a
tubular calyx and the blades spreading ; and the

Cruciferous, of four somewhat similar petals, the four abruptly
spreading blades in the form of a cross (erucrate), as in Fig. 394.

Rosaceous, with roundish and widely spreading petals on very
short or hardly any claws, as in Rose and Apple-blossoms.

Liliaceous, a 6-phyllous perianth of campanulate or funnelform
shape ; the members either distinct, asin most common lilies and
tulips, or gamophyllous, as in Lily of the Valley. All but the
first and last of these sorts are examples of regular and chori-
petalous perianth.

Orchidaceous flowers are of a peculiar irregularity, combining
both calyx and corolla: one member, the petal in front of the

476

FIG. 475. Rotate or wheel-shaped and five-parted corolla of the Bittersweet (So-
lanum Duleamara).

FIG. 476. Whiecl-sheped and five-lobed corolla of the common Potato.

PERIANTH, OR FLOWER-LEAVES. 247

stamen and stigma, differs from the rest in shape and in being
nectariferous (as in Fig. 460) ; it is named the Lapertum.

Galeate is a term applied to a corolla the upper petal or part
of which is arched 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 latter, it
consists of two, completely united.

450. Gamophyllous forms with special names are chiefly the
following. Illustrations are usually taken from the corolla, but
the forms and terms are not peculiar to it, excepting the first,
viz. the

Ligulate or Strap-shaped corolla (Fig. 288, &c.), which is
nearly confined to Compositz. Here a corolla, formed of three
or five petals, imitates a single petal, except at its very base,
which is commonly tubular: the remainder is as though the tube
had been split down on the upper side and flattened out. The
corolla of Lobelia, type of a family most nearly related to Com-
posite, illustrates this. (Fig. 488.)

4i7 478 479 480 481
451. The names of the general forms are mostly taken froim
some resemblance to common objects. All those in common
use will be found in the Glossary: a few leading ones are here
specified. They may be divided into the regular and the irregu-

lar. The principal irregular form with a special name is the
Labiate, or lipped, also 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. This bilabiate character in the corolla, and often in
the calyx also, pervades several orders with gamopetalous
flowers, and gives name to one of them, the Labiatz, to which

FIG. 477. Campanulate corolla of the Harebell, Campanula rotundifolia. 478.
Salverform (hypocraterimorphous) corolla of Phlox. 479. Labiate (ringent) corolla of
fLamium; a side view 480. Personate corolla of Antirrhinum or Snapdragon. 481.
Personate corolla of Linaria, spurred (calearate) at the base.

248 THE FLOWER.

the Sage and Mint belong. Such flowers are 5-merous, and
have two members specially united to form one lip, and three in
the other. The odd sepal being posterior (or next the axis of
inflorescence) , and consequently the odd petal anterior, the calyx
has its lower lip of two sepals and its upper of three; while the
corolla has its upper lip of two petals and its lower of three.
But in Leguminose, where the calyx is sometimes bilabiate, and
where the odd sepal is anterior (or toward the bract), this is
reversed, and two sepals or lobes of the calyx form the upper
lip and three the lower. A bilabiate corolla is

Ringent, that is gaping or open-mouthed, when the throat is
freely open, as in Lamium, Fig. 479 ;

Personate, or masked, when the throat is closed, more or less,
by a projection of the lower lip called the PaLatr, as in Antir-
rhinum and Linaria, Fig. 480, 481.

452. Of regular forms, there are the following, beginning with
that having least tube:

Rotate, or Wheel-shaped (Fig. 475, 476), widely spreading from
the very base, or from a short and inconspicuous tube.

Crateriform, or Saucer-shaped, like rotate except that the broad
limb is cupped by some upturning toward the margin.

Hypocrateriform, or rather (not to mix Latin and Greek)
Hypocraterimorphous, in English Salverform, when a rotate or
saucer-shaped limb is raised on a
slender tube which does not much
enlarge upward ; that is, where a long
and narrow tube abruptly expands
into a flat or flattish limb, as in
Fig. 478. In Fig. 472-474 are seen
salverform corollas with somewhat
more upwardly dilated (trumpet-
shaped) tube. The salver or hypv-
eraterium, which the name refers to,
with a stem or handle beneath, is now
to be met with only in old pictures.

Tubular, when strictly used, denotes
a gamophyllous perianth with limb
inconspicuous in proportion to the
tube, as in Trumpet Honeysuckle, or as Fig. 472-474 would be
if the limb were much diminished or wanting. But it is some-
times used in the sense of having a conspicuous tube.

FIG. 482. Calyx and funnelform (infundibuliform) corolla of a common Morning-
Glory, Ipomcea purpurea.

ANDRGCIUM, OR STAMENS. 249

Infundibuliform, or Funnelform, such as the corolla of common
Morning-Glory (Fig. 482), denotes a tube gradually enlarging
upward from a narrow base into an expanding border or limb.

Campanulate, or Lell-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. THE ANDRG@CIUM, OR STAMENS IN PARTICULAR.

453. The whole Stamen. For the general character and some
of the modifications of the stamens, see the first (501) and por-
tions of the succeeding sections of the present chapter. The
terms peculiar to these organs, and of common use in botanical
description, were nearly all coined by Linnzeus, and employed as
the names of classes in his sexual system. (672.) The sub-
stantive names of those classes which are characterized by the
number of stamens, and which were designated by Greek nume-
rals prefixed to andria (the Greek word for man being used
metaphorically for stamen), are put into adjective form, as
follows :

Monandrous, for a flower with a solitary stamen ; Diandrous,
for a flower with two stamens; Z7riandrous, with three; Tetran-
drous, with four; Pentandrous, with five ; Hexandrous, with six ;
Heptandrous, with seven; Octandrous, with eight ; Lnneandrous,
with nine; Decandrous, with ten; Dodecandrous, with twelve;

Polyandrous, with a greater or indefinite number, or /eosandrous
(meaning twenty-stamened) when a polyandrous flower has the
stamens inserted on the calyx, as in the Cherry (Fig. 337),
Pear, &c.

FIG. 483. Diadelphous stamens (9 and 1) of a Pea. 484. Monadelphous stamens of
a Lupine. 485. Monadelphous stamens, &c., of Mallow.
FIG. 486. Five syngenesious stamens of a Composita. 487. The same, laid open.

250 THE FLOWER.

Didynamous is a term applied to an andreecium of four sta-
mens in two pairs, a longer and a shorter, as in Fig. 361.

Tetradynamous is similarly applied to that of six stamens, two of
them shorter, in the manner characteristic of Cruciferz, Fig. 396.

454. Terms which denote coalescence of stamens, whether by
their filaments or their anthers, are

Monadelphous, that is, in one brotherhood, by coalescence of
the filaments into a tube, as in the Mallow (Fig. 485), Lupine
(Fig. 484), Lobelia (Fig. 488), &e.

Diadelphous, in two brotherhoods, by coalescence of the fila-
ments into two sets; sometimes an equal number in each, as in
Fumariacee (Fig. 390), sometimes nine in one set and one
separate, as in the Pea (Fig. 483) and most Papilionaceze.

Triade/phous, with filaments united in three sets or clusters, as
in Hypericum.

Pentadelphous, in five sets, as in Linden, Fig. 398, 399.
But in general, when the sets are several, without regard to the
number the stamens are said to be Polyadelphous.

Syngenesious, when the stamens are united by their anthers
into a tube or ring; as in the whole vast order of Composite

489 491 492

(Fig. 486, where they are five in number and the filaments dis-
tinct), in Cucurbita (Fig. 489, 490, where they are three in
number and the filaments partly monadelphous), and in Lobelia
(Fig. 488, where they are also five and the long filaments are
mainly monadelphous).

FIG. 488. Flower of Lobelia cardinalis, with tube of corolla divided on one side;
filaments and anthers united into a tube: f. tube of filaments; a. of anthers.

FIG. 489. Male flower of Cucurbita (Squash), with limb of calyx and coroHa cut
away, to show the stamens, viz., three filaments, separate at base but monadelphous
above, and three syngenesious anthers ina kind of head. 490. Stamens of the same,
enlarged and the upper part cut away, to show the union. The anthers are sinuous.
491. A detached stamen of the Melon, with loosely sinuous anther. ;

Fig. 492. Stamens and style of a Cypripedium, united into one body or co/umn:
@, anthers; st. enlarged sterile stamen; stig. the stigma.

ANDRG&CIUM, OR STAMENS. 251

455. Of terms relating to adnation of stamens, besides the
general ones of hypogynous, perigynous, epigynous (332), and
epipetalous, or adnate with corolla, there is the special one of

Gynandrous, having stamens borne upon the pistil, as in
Orchidacex. 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 proper Orchis
tribe (as in Fig. 460, 461), anther and stigma are consolidated
into one mass, and there is no evident style.

456. A complete stamen consists of Fmamenr and ANTHER.
The latter is the functionally essential part of the organ, and
therefore is wanting only in abortive or sterile stamens. (345,
352, &c.) The filament, being only a stalk or support, may be
very short or wholly wanting: then the anther is sessile, just as
the blade of a leaf’ is said to be sessile when there is no petiole.

457. The Filament, although usually slender and stalk-like,
assumes a great variety of forms: it is sometimes dilated so as
to resemble a petal, except by its bearing an anther; as in the
transition states between the true petals and stamens of Nym-
phzea, 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 appendaged by outgrowths. Some
of these appendages are very 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 species of Allium.

459. The Anther, the essential organ of the stamen, contain-
ing the pollen, surmounts 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 rarely
divided into two cavities (locelli), the best technical name for
anther-sac is that of THeca. The two theese, lobes, or cells are
commonly connected by a more or less evident and sometimes
conspicuous common base or junction, which is mostly a pro-
longation of the filament, the Connectrvum, or in English
CONNECTIVE.

460. For the discharge of the pollen, the cells of a normal
anther open at the proper time by a line or chink, usually

252 THE FLOWER.

extending from top to bottom (Fig. 493), the suture or line of
dehiscence. Commonly this line is lateral or marginal: not
rarely it faces forward or backward. In the vast genus Solanum,

i)

495 44. 495 45 497 * 498

to which the Potato belongs, in most Ericaceous plants (Fig.

458, 494), in Polygala, and in many other flowers, the anther-

cells open only by a hole (foramen or pore), or at most a short

chink, at the tip, through which the pollen has in

some way to be discharged. In Vaccinium (Cran-

berry, Blueberry, &c.), the pore-bearing tip of the

anther-cell is prolonged considerably, often into a

slender tube, as in Fig. 340. In the Barberry (Fig.

495) and in most of that family, also in Lauracez,

~™ the whole face of each anther-cell separates by a con-

tinuous line, forming a kind of door, which is attached at the top,

and turns back, as if on a hinge: in this case, the anthers are said

to open by uplifted valves. In the Sassafras and many other

plants of the Laurel family, each lobe of the anther opens by two
smaller valves of the kind, like trap-doors.

461. The attachment of the anther to the filament presents
three principal modes, which are connected by gradations.
These are the

Innate (Fig. 495, 496), in which the anther directly continues
and corresponds to the apex of the filament, the cells usually
dehiscent strictly marginally, the lobes or cells not looking or
projecting either inward or outward.

FIG. 493. A stamen, with its anther, 6, surmounting the filament, a, and opening
in the normal manner down the whole length of the outer side of each cell.

FIG, 494. Stamen of a Pyrola; each cell of the anther opening by a terminal ori-
fice or pore.

FIG. 495. Stamen of a Barberry; the cells of the anther opening each by an up-
lifted valve.

FIG. 496. A stamen of Isopyrum biternatum, with innate anther. 497. Stamen of
Liriodendron, or Tulip-tree, with adnate extrorse anther. 498. Stamen of CEnothera
glauca, with the anther fixed hy its middle and versatile

FIG. 499. A stamen of Asarum Canadense, with adnate anther and prolonged tip
to connective.

ANDRGECIUM, OR STAMENS. 253

Adnate, in which the connective appears to be a direct con-
tinuation of the filament, having the anther adherent to the
anterior or posterior face of it, and the lines of dehiscence
therefore looking inward or outward. Magnolia, Limodendron
(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.

462. The direction to which an anther faces, whether inward
{toward the centre of the flower), or outward (toward the peri-
anth), has to be considered; except in the case of an innate
anther with strictly lateral or marginal dehiscence. An anther is

Extrorse, i.e. turned outward, or Posticous, when it faces to-
ward the perianth, as in Magnolia and Liriodendron (Fig. 497),
Asarum (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.

Introrse, i. e. tarned inward, or Anticous, when it faces toward
the axis of the flower; as in Nymphzeacee (Fig. 318), in Violet
and Lobelia (which are adnate and introrse), and in Génothere.
In the common Evening Primroses (as in Fig. 498) the anther
is fixed near the middle, introrse, and versatile.

463. The direction in which the anther may be said to face,
outward or inward, depends 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 strongly
characterized adnate anther as that of Liriodendron (Fig. 497),
both the attachment and the dehiscence are plainly posticous or
extrorse: in most species 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 extrorsely affixed toward the base
or middle to a slender tip of the filament, the dehiscence is
either introrse or nearly marginal. Parnassia is in similar case ;
the anthers being clearly extrorse as to insertion and more or
less introrse as to dehiscence.

464. Adnate anthers are perhaps as frequently extrorse as
introrse. Others, whether bas/fixed or med/ fired, are more com-
monly introrse. ‘Those fixed by the middle, or at any other
part of the back, and lying on the inner side of the filament,
are said to be Incumbent.

954 THE FLOWER.

465. The connective may be appendaged either by a prolon-
gation or otherwise from the tip (as in Fig. 499), or from the
back, as in Violets and in many Ericaceous plants.

466. The normal anther is two-celled, bilocular, or (to use
a less common term) didiecous, and its lobes or cells parallel,
right and left; but the cells at first, and sometimes at maturity,
are bilocellate, that is each is divided into two by a partition
which stretches from the connective to the suture or line of
dehiscence. In an
innate anther, and
B | in many others, this
Y \ine of dehiscence is

marginal or lateral,

either stricily or
a ne =e nearly so, as in
Fig. 500. When introrse or extrorse (as in Fig. 501, 502),
the sutures may still be considered to represent the margins
turned inward or outward. The pollen is accordingly pro-
duced in four cavities or separate portions of the interior. But
the two locelli on the same side of the midrib or connective
(right and left) are usually confluent into one pollen-filled cavity
or cell at maturity if not earlier, or at least the partition between
them breaks up at dehiscence. Sometimes it remains, and, the
groove at the sutures being deep, the anther is strongly four-
lobed or quadrilocular at maturity, as in Menispermum (Fig.
504) ; but morphologically this is still only bilocular (dithecous)
although quadrilocellate, and the anther opens at the sutures
and through these partitions.

467. A stamen being the homologue of a leaf, the natural
supposition is that the anther 1s homologous with the blade or
an apical portion of the blade, therefore the two lobes or thecz
with the right and left halves of it, the intervening connective
with the midrib, and the line of dehiscence with the leaf-mar-
gins.1 This conception is exemplified by the accompanying

1 This is the view long ago taken by Cassini and Reeper, and it may still
be maintained as the best morphological conception. Mohl interposed some
objections to its universality ; but, as presented in Sachs’s Text-Book, they
are not incompatible with the common morphology. Sachs takes the fila-
ment with the connective to be the homologue of the whole leaf, and the
anther-cells as appendages. Others, in likening the anthers to glands, adopt
a similar view.

FIG. 500. Innate anther, same as Fig. 496. in younger state, with transverse section,
showing the four locelli. 501. Same of an adnate extrorse anther, such as Fig. 497.
502. Same as the preceding but mature and dehiscent, the two locelli becoming one cell
hy the vanishing or breaking up of the partition.

ANDRGECIUM, OR STAMENS. 255

diagram, Fig. 503, which should, however, show the median
partitions in the cross-section, or traces of them. Pollen is a
special development into peculiar cells of what would
be parenchyma 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 early be confluent into one cell.

468. Of the many deviations of the typical 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 carly
development, become confluent at the apex, as in the 5s
Mallow family (Fig. 506), 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 way, 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.

469. 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 Monarda (Fig.
508), in which their contiguous ends
so coalesce as to give the appearance
of a one-celled anther fixed by the
middle. Or, again, the two cells may
be separated by the enlargement of
the connective between them, asin 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

oH 50D 506 HOT

FIG, 503. Diagram to illustrate the morphology of the stamen, on the idea that the
anther answers to leaf-blade: the lower portion being filament and a] art of the anther,
in section, the upper a part of a leaf.

FIG. 504. Stamen of Menispermum Canadense, the quadrilocellate anther divided.

FIG. 505. Stamen of Pentstemon pubescens, with anther-cells divergent.

FIG. 506. Stamen of Mallow (one of the cluster of Fig. 485), the two cells and sutures
confluent into one.

FIG, 507. Anther of Gomphrena or Globe Amaranth, medifixed, of a single cell,
dehiscent.

256 THE FLOWER.

proper filament, and bearing an anther-cell at each end. Ina
few species, the two anther-cells are nearly alike; in more, the
lower one is imperfect, as in Fig. 510%; in more, it is abortive
or wanting altogether, as in Fig. 510°. Then, in the related
Californian genus Audibertia, the lower half of this connective

3S 50D

is reduced to a short tail, as shown in Fig. 511%, or even in
most of the species to so minute a vestige that, except for these
transitions, the stamen might be supposed to consist of a simple
filament, with an interruption like a splice in the middle, and
surmounted by a one-celled anther, as shown in Fig. 511°. In
Rosemary, the continuity is complete, although a minute reflexed
tooth sometimes indicates the junction.

470. Pollen, the product of the anther, is usually a powdery
substance, which when magnified is seen to consist of separate
grains, of definite size and shape, uniform in the same plant,

512 515 54 51S
but often very different in different species or families. The
grains are commonly single cells, globular or oval in shape, and
of a yellow color. But in Spiderwort they are oblong; in the

FIG. 508-511. Anthers, with upper part of filament. of several Labiate. 508. Of
Monarda. 509. Ofa Calamintha. 510 Of two species of Salvia. with long and slender
connective, the upper fork of which bears one anther-cell; the lower in @ (from Salvia
Texana), bearing the othercell in an imperfect condition; in b (from S. coccinea), bear-
ing none at all. 511. a. Same of Andibertia grandiflora, the lower fork of the connec-
tive reduced to a naked spur; b, from A. stachyoides, in which this lower fork is nearly
wanting, and the upper is in a straight line with the filament which it seems to
continue.

FIG. 512-515. Forms of pollen: 512, from Mimulus moschatus; 513, Sicyos; 514.
Echinocystis; 515, Hibiscus.

POLLEN. eg

Cichory and Thistle tribes, many-sided; in the Musk-plant,
spirally grooved; in the Mallow family and the Squash and

Pumpkin, beset with briskly projections, &c. The pollen of
Pine, as well as that of the Onagracez, is not so simple, but
appears to consist of three or four blended cells ;
that of most Ericaceze 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 they lie side by side in the anther,
resemble a skein of silk.

471. Pollen-grains are usually 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 shreds, forming the cobweb-like
threads mixed with the pollen of Evening Primrose,
or as a kind of tissue combining the pollen into
coherent masses, of various consistence. Of this
kind are the elastically coherent pollen-masses (or
Poti, sing. PoLirytum) of Orchises (Fig. 463),
and the 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 fleshy. This is later
formed than the inner, and by a kind of secretion from it: to it
all the markings belong. The inner coat, which is the proper
cell-wall, is a very thin, delicate, transparent and colorless mem-
brane, of considerable strength for its thickness. The pollen
of Zostera 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 slightly turbid under the higher
powers of ordinary microscopes, and, when submitted to a mag-

FIG. 516-521. Forms of pollen: 516, Lily; 517, Cichory; 518, Pine; 519, Circea;
520. Kalmia; 521, Evening Primrose.
FIG, 522. <A pair of pollinia of Asclepias. annexed by their caudicles to the gland.

17

258 THE FLOWER.

nifying power of about three hundred diameters, is found to
contain a multitude of minute particles (foville), the larger of
which are from one four-thousandth to one five-thousandth of an
inch in length, and the smaller only one fourth or one sixth of
this size. When wetted, the grains of pollen promptly imbibe
water by endosmosis, and are distended, changing their shape
somewhat, and obliterating the longitudinal folds, one or more
in number, which many grains exhibit in the dry state. Soon
the more extensible and elastic inner coat inclines to force its
way through the weaker parts of the outer, especially at one or
more thin points or pores; sometimes forming projections,
when the absorption is slow and the exterior coating tough. In
many kinds of pollen, the grains, when immersed in water, soon
distend to bursting, discharging the contents.

474. Pollen-tubes. In others, and in most fresh pollen, when
placed in ordinarily aerated water, at least when this is slightly
thickened by syrup or the like, and submitted to a congenial tem-
perature, a projection of the inner coat through the outer appears
at some one point, and by a kind of germination grows into a
slender tube, which may even attain two or three hundred times
the diameter of the grain; and the richer protoplasmic contents
tend to accumulate at the farther and somewhat enlarging ex-
tremity of this pollen-tube.?

475. In cleistogamous 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
directing itself toward the stigma in a wonderful way. Similarly,
in the open flower of Milkweeds, the pollen-tubes sometimes
start from the pollen-mass even while yet in the anther, and in
vast numbers, forming a tuft or skein of pollen-tubes, which
may attain considerable length and direct itself toward the some-
what distant stigma. Commonly, however, the pollen remains

1 In Conifer, the grains of pollen have a peculiar internal structure or
rather a development (suggestive of a homology with the microspores of some
of the higher Cryptogamia), the contents at or before maturity undergoing
division into two or three internal cells, only one of which acts in fertiliza-
tion. When they act upon the ovule or are placed in water, and the inner
coat swells by absorption, the bursting outer coat is commonly thrown off.
In Pines and Firs (but not in Larch and Hemlock Spruce), the grain of
pollen is singularly compound, consisting (as in Fig. 518) of a central arcuate
body (the proper pollen-cell) bearing at each end an empty roundish cell.
These are vesicular protrusions or appendages of the proper pollen-grain, of
no known functional importance, except that they render such wind-dis-
persed pollen more buoyant for transportation.

2 Van Tieghem, in Ann. Sci. Nat. ser. 5, xii. 812, &e., 1869.

ay

GYNCECiIUM IN ANGIOSPERMS. 259

unaltered until it is placed upon the stigma. The more or less
viscid moisture of this incites a sim- = oe 2
ilar growth, and also doubtless nour- A
ishes it; and the protruding tube at
once penetrates the stigma, and by glid-
ing 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 by the action of this
pollen-tube upon the oyule, and upon a
special formation within it. Consequent 526

upon this an embryo is formed ; and the ovule now becomes a seed.

Section VII. Tue Pistits, or Gynacium.
§ 1. Ix 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 gyneecium in Section III., relates to the most
typical conditions of this part of the flower. The essential
characteristics of all ordinary pistils, whether simple or compound,
are: 1. a closed ovary, in which one or more ovules are included ;
and 2. a stigma, upon which pollen for fertilizing the oyules is
received, and through which the pollen acts upon them. There
is a more simplified condition, in Gymnosperms, 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 seeds enclosed in a sac or covering, this
in the flower being the ovary.’ And plants with such gynecium
are denominated ANGIOSPERMS or ANGIOSPERMOUS PLANTS. ‘To
such only the present subsection specifically relates.

477. The several terms which apply to the Gyneetum or
female system of a flower, and to its components, have been

1 Although thus originated, the seeds are not in all cases matured in a
closed pistil. In the Blue Cohosh, Caulophyllum thalictroides, the ovules
rupture the ovary soon after flowering, and the seeds become naked; and in
Mignonette they are imperfectly enclosed, the ovary being open at the
summit from an early period of fructification.

FIG. 523. <A pollen-grain of Datura Stramonium, emitting its tube. 524. Pollen-
grain of a Convolvulus, with its tube. 525. Other pollen-grains, with their tubes, less
strongly magnified. 526. A pollen-grain of the Evening Primrose, resting on a portion

of the stigma, into which the tube emitted from one of the angles penetrates; the oppo-
site angle also emitting a pollen-tube. All highly magnified.

260 THE FLOWER.

enumerated and defined already (302, note): the elementary
term is that of

478. Carpel, Lat. CarpeLttuM. This is the term coined by Dunal,
and is in common use. The better-formed word Carprprum
(English Carpid) has been proposed, and best of all Carpo-
PHYLLUM, in English Carpopiyll. For carpels are, as the word
carpophylla denotes, pistil-leaves, or leaves of the gyncecium,
7. e., seed-bearing or fructiferous phylla. They occupy the cen-
tral or uppermost region of the flower. A carpel may be a pistil
of itself, either the only one of a blossom or one of several, or
it may be a constituent of a more complex pistil. In either case,
a carpel is the homologue of a leaf.

479. The morphological conception of an uncombined carpel
is that of the blade of a leaf incurved lengthwise, so that
the margins meet, and join by a suture, thus forming a closed
sac, the ovary. A prolongation of the tip of the leaf is the style:
some portion of this, usually the apex, not rarely a single or
double line down the side which answers to the suture of the
leaf-margins, and may be regarded as its continuation, is the
stigma. ‘The carpellary leaf is always /zcurved: the lower sur-
face of the leaf is represented by
the exterior surface of the ovary,
the upper by the interior. The
conjoined margins of the leaf, or
whatever they bear, are internal
in the ovary: the stigma may be
regarded as a portion of leaf-
margins presented externally, des-
Y titute of epidermis and formed
be pot = of loose cellular tissue, which in
anthesis is moist by some secretion. The ovwes are peculiar
structures normally arising as outgrowths from the margins of
the leaf, or some part of them, sometimes from the whole or
a special portion of the upper or inner surface of the leaf.

480. The carpellary leaf being involute, the suture, on which
the ovules are normally borne, always looks toward the axis or
centre of the flower. It is the only proper suture (or seam) a
carpel can have. From its position it takes the name of /ener
or Ventral Suture. And the opposite line or ridge, answering
to the midrib of the leaf, being sometimes prominent and of the

FIG. 527. A leaf incurving, to illustrate the morphology of a simple pistil or carpel.
528. A carpel (of Isopyrum biternatum), cut across, the lateral stigma (here manifestly
a double line) and the suture bearing the ovules turned toward the eye. 529. A ripe
carpel of Marsh Marigold which has opened and shed the seeds: the points of attach-
ment of the latter conspicuous along the edges of the carpel.

GYNCCIUM IN ANGIOSPERMS. 261
appearance of a suture, has been somewhat incongruously named
the Outer or Dorsal Suture.

481. The number of carpels in a gyneecium is simply expressed
by adjective terms consisting of Greek numerals prefixed to this
word: e. g., Monocarpellary, of a solitary carpel ; Dicarpellary, of
two carpels; Zricarpellary, 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 only partially synonymous terms are such as Monogynons
(of one pistil), Digynous (of two), Polygynous (of many), &e.
These are adjective forms of the names of the orders, from
Monogynia to Polygynia, in the Linnzean artificial classification,
which either supposes the carpels to be separate or partly so, or
confounds simple and compound pistils.

482. When the gyneecium is of a solitary 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 oyule-bearing suture
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 usually median, that is antero-posterior or in the
line of bract and axis. Cruciferse, Capparidacez, and Fumari-
acee are somewhat remarkable for having their two carpels
right and left, that is, collateral or, in other words, transverse.
When three, four, or a greater number, they divide the circle
equally, or when numerous they take a spiral instead of verticil-
late order, and occupy several or many ranks, as in Ranunculus,
Magnolia, Potentilla, &c.

483. The Gynecium may be either of separate carpels
(Apocarpous), or of carpels coalescent into one body (Syncar-
pous), or of all grades between the two. Apocarpous pistils are
simple ; a syncarpous pistil is compound.

484. In both, the essential parts are the ovary and the stigma.
The style may be conspicuous and widely separate these two,
as in Fig. 536-538 ; or hardly 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
interior of the ovary on which ovules are borne. It has been
stated (579) that these are usually borne upon the margins of

1 Taken from aremote analogy with the placenta of the higher animals.
The name appears to have been introduced into botany by Adanson. It has
been termed Trophospermum or Spermophorum by some of the early modern
botanists.

262 THE FLOWER.

ihe carpellary leaf, or upon some portion of what answers to
them. When the ovules are numerous, and some-
times when they are few, the combined leaf-edges
enlarge to form a kind of receptacle for their attach-
ment or support: this is the Placenta. In Fig. 530,
the placenta is well developed, and also in such
syncarpous ovaries as are illustrated in Fig. 536,
537, 544, and 545. In very many others (such as
Fig. 528, 531, 533), there is no particular enlarge-
ment of the leaf-margins visible, and no particular
ground for the use of this special term. Still it
is commonly used, as occasion serves, even for the
mere line or spot on which ovules are borne, as
aie as for a more prominent development to which
530 the name was originally applied.

486. Simple or Apocarpous Pistils may be solitary, several, or
numerous. When indefinitely numerous, they are seldom in one
circle, but are capitate or spicate upon a proportionately enlarged
or prolonged receptacle, as in Anemone, Ranunculus, and most
strikingly in Myosurus; when reduced to a single one, as in
Acta, Podophyllum,* Barberry, and Plum or Cherry, the car-
pel mostly appears as if it were an actual termination of the
floral axis. But even then the pistil
is hardly ever quite symmetrical in
shape: the ovary is somewhat gib-
bous or unequal-sided (as in Fig,
312,315, 316, 528, 531-533), and the
stigma more or less oblique or even
wholly lateral. The continuation of

331 532 533 the latter down the whole length of
the ventral side of the style (as in Fig. 528, and also Fig. 549)
is not uncommon. In Schizandra (Fig. 531) it is continued
downward on the ventral edge of the ovary as far as to its
middle.*

1 Abnormal specimens of Podophyllum peltatum are occasionally found
having a gynecium of from two to six separate carpels.

2 Pleurogyne, a Gentianaceous genus so named on this account, has no
style nor apical stigma whatever, but has along stigma extending down the
outside of each ovuliferous suture of its dicarpellary ovary for most of its
length.

FIG. 530. Single simple pistil of Podophyllum, cut across to show the placenta, &c.

FIG. 531. Vertical section of a pistil of Schizandra coccinea; a side view showing
the stigma decurrent down to the middle of the ovary. 532. Pistilof Hydrastis; ventral

view. 533. Pistil of Actsea rubra, cut across, so as to show the interior of the ovary;
ventral view.

GYNGECIUM IN ANGIOSPERMS. 263

487. As the placenta of a simple pistil belongs to the two
united margins of the carpellary leaf, there is naturaily a double
row of ovules, one to each margin. If the leaf-
margins which are turned inward in the ovary be-
low to bear the ovules are turned outward above to
receive the polien (see Fig. 531), then the typical
stigma should also be double or bilamellar. So it
is seen to be in such carpels as those of Fig. 528,
531-533, and indeed in very many stigmas of this
class. Such division, or even a greater bifurcation
of a monocarpellary stigma into two lobes or half-
stigmas, is not anomalous.

488. The ovary of a simple pistil should be
untlocular, 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 bilocellate, 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. Compound or Syncarpous Pistil.’| This consists of two.
three, or a greater number of carpels coalescent into one body.
A true compound pistil represents a whorl (in the simplest case
a pair) of carpels united into one body, at least as to the ovary.

490. The coalescence of a capitate or spicate mass of carpels
or simple pistils of the same flower, imbricately heaped on the
torus, as in Magnolia (Fig. 648) and Liriodendron, cannot
properly be said to form a compound pistil. This heap of
pistils may be called a Sorema.

491. Morphologically, a compound pistil, as to the ovary, may
be a pair or a circle of closed carpels or simple pistils brought
into contact, and the contiguous parts united: this is illustrated,
in Fig. 535-538. Or it may be formed of a whorl of open car-
pellary leaves, joined each to each by the contiguous margins,

5o4

1 The terms apocarpous and syncarpous for pistils, the first of separate, the
second of combined carpels, were introduced by Lindley. They have little
advantage over the terms simple and compound. Moreover, the word
syncarp or syncarpium had been appropriated to a sort of fruit of the class
now called 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. 534. Ovary or forming legume of Astragalus Canadensis, transversely divided,

to show the false partition which, intruded from the back, divides the simple cell into
two half-cells or locedii,

264 THE FLOWER.

in the manner of Fig. 542-545. Between these two there is
every gradation. ‘The first forms a compound ovary,

492. With two or more Cells and Axile Placente. For it is
evident that, if the contiguous parts of a whorl of two or more
closed carpels cohere, the resulting compound ovary should have
as many cells as there are carpels in its composition, and that
the placentze (one in the inner angle of each carpel) will all be

585

brought together in the axis of the compound pistil. And the
partitions, termed Dissrpmrents, which divide the compound
ovary into cells, manifestly consist of the united contiguous por-
tions of the walls of the carpels. These necessarily are composed
of two layers, one belonging to each carpel; and in fruit they
often split into the two layers. True dissepiments and the true
cells must accordingly be equal in number to the carpels of
which the compound pistil is composed. That is, the ovary, or
the resulting fruit, is bélocular or 2-celled, trilocular or 3-celled,
quadrilocular or 4-celled, and so on, according to the number of
dissepiments or cells.

493. There may also be false dissepiments, mostly of the same
character as that which in Fig. 534 divides the cell of a singie
carpel. Such are found in Flax (Fig. 539-641), in Amelanchier
or Service-berry, in Huckleberry (Gaylussacia), and in most of

FIG. 535. Pistil of a Saxifrage composed of two carpels or simple pistils united
below, but distinct above; cut across both above and below.

FIG. 536. Pistil of common St. Johnswort, of three united ovaries; their styles
distinct.

FIG. 537. The same of another species of St. Johnswort (Hypericum prolificum),
the styles also united into one, which, however, may split apart in the fruit.

FIG. 538. Pistil of Tradescantia or Spiderwort, even the three stigmas united into
one. The ovary in all cut across to show the internal structure.

vet ie

GYNCECIUM IN ANGIOSPERMS. 265

the American species of Vaccinium. In all these, the false par-
tition is a growth from the middle of the back of each carpel,
which divides its cell more or less completely into two.

on) 540 Hl

494. On the other hand, even the true dissepiments which
belong to such a compound ovary may be abortive or evanescent,
the placentze remaining in the axis combined into a column.
(499.) The second modification of the compound pistil (491)
normally has an ovary,

495. With one Cell and Parietal Placente. That is, the
placentz are borne (as the term denotes) on the wall or parietes
of the ovary, as in the Poppy, Violet, Sundew,
Cistus or Helianthemum (Fig. 543), Cleome, Gen-
tian, and in all or most of the orders from which
these examples are cited. The diagram Fig. 542
illustrates the morphological conception of a com-
pound pistil of this kind. Not that it is ever sup-
posed to be formed by the actual combination of once

544 545

separate leaves, any more than a gamophyllous calyx or corolla
is actually so produced. The conception in all such cases is that

FIG. 539. Transverse diagrammatic section of a flower of the common Flax, show-
ing the ovary with false partitions extending one from the back of each cell. 540. Sec-
tion of a mature fruit and seeds of the same, the false partitions now complete, divid-
ing the five cells into ten, each one-seeded. 541. Same of a wild Flax (Linum perenne),
in which the false partitions remain incomplete.

FIG. 542. Plan of a one-celled ovary with three parietal placenta. cut across be-
low; the upper part showing the top of the three leaves it is theoretically composed of,
approaching, but not united.

FIG. 543. Ovary of Helianthemum Canadense, cut across, showing the ovules on
three parietal placente.

FIG. 544. Transverse section of the ovary of Hypericvm graveolens; the three large
placenta meeting in the centre, but not cohering. 545. Similar section of a ripe capsule
of the same; the placentze now evidently parietal

256 THE FLOWER.

of a congenital development of organs in union which, in the
development of a vegetative shoot, would be leaves.* This case
is represented by the combination of open carpellary leaves, as
the preceding one is by that of closed ones. As the edges
of the leaves must needs be turned in, to bear the ovules,
a compound ovary with parietal placentation may be likened
to the unopened calyx of a Clematis, as shown in Fig. 256,
257. Every gradation is found between axile and parietal
placentation. Sometimes the placentz are strictly on the pari-
etes or wall (Fig. 545, 547); sometimes borne inwards on
incomplete dissepiments (Fig. 548); and sometimes they are
brought firmly together in the axis, as in Fig. 544, though sepa-
rable, and indeed separated in the fruiting stage. _

496. A compound ovary with parietal placentz is necessarily
one-celled (unilucular) ; except it be divided by an anomalous
partition, such as is found in Cruciferz (Fig. 395) and in many
Bignoniacee.

497. Normal placentz are necessarily double: when parietal,
the two halves belong to different leaves ; when axile, to the same
leaf. These two halves may diverge or be widely separated,

sometimes even at their origin, as in Aphyllon and some other
Orobanchacez, in which a dicarpellary ovary has four almost
equidistant placentze; or in such cases the placentz may be
regarded as intra-marginal instead of marginal.

498. The placentz of a two-several-celled ovary, such as in
Fig. 536, 537, &e., may be described in the plural number,
being one in each carpel: or when consolidated into a central
column, and well covered with ovules, they may be said to form
one (compound) placenta. Then when the dissepiments early
disappear, or are abortive from the first, the result is a compound
ovary of this class,

499. With one Cell and Free Central Placenta. In Caryo-
phyllacee (Fig. 549, 550) and Portulacacez, this evidently
results from the obliteration of the dissepiments (as many as
there are styles or stigmas), vestiges of which may be sometimes

FIG. 546. Diagram (ground-plan) to illustrate free central placentation produced
by abortion of dissepiments. 547. Same of strict parietal placentation. 548. Same with
the placentz carried inward on imperfect dissepiments.

GYNGCIUM IN ANGIOSPETMS. 267

detected, while certain plants of the same familics, of otherwise
identical structure, retain the dissepiments even in the fruit.

500. But a similar condition may equally arise
from a modification of parietal placentation, namely,

with the margins of the leaves ovuliferous only at \
p

549

bottom, and the placentze there conspicuously devel-
oped and completely united. The basal placenta-
tion of Dionzea is unavoidably so explained, its
nearest relative, Drosera (Fig. 553), having parietal
placentze. And this leads to a probable explanation
of the case in Primulacez, where a large free central
placenta fills the centre of the cell, and no trace of
dissepiments can be detected.’

501. The idca maintained in former editions is
still adhered to; namely, that placentz belong to
cearpels and not to the cauline axis, in other words,
that ovules are productions of and borne upon leaves, usually
upon their margins, not very rarely upon other portions of their
upper surface, rarely 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 Nym-
pheea and Nuphar, Fig. 551) excepting the inner angle, to which
they are usually restricted in other plants. And in the allied
Brasenia and Cabomba, where the ovules 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-
growth from the base of the carpellary leaves, combined over the floral
axis. Upon this interpretation, 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. 529 (1869) ;
Celakowsky, Vergleichende Darstellung der Placenten, &c. (1876) ; Wariing,
in Ann. Sci. Nat. ser. 6, v. 192.

2 This view was first maintained as a general theory, and on critical
grounds, by Brown, in Plante Javanice Rariores, 107-112. Schleiden, End-
licher, and others took the opposite view, 7. 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 espe-
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 l’Ovule, in Ann.
Sci. Nat. ser. 6, v. 1877-78.

FIG. 549. Vertical section through the compound tricarpellary ovary of Spergularia
rubra, showing the free central placenta. 550. Transverse section of the same,

268 THE FLOWER.

margins of the carpellary leaf. In many species of Gentian, as
also in Obolaria and Bartonia, of the same
family, the whole internal face of a dicar-
pellary ovary is thickly ovuliferous.
503. Perhaps the parietal placentae in
Parnassia (Fig. 552) are borne on the
midribs of the carpels, for they are directly
= * under the stigmas, instead of alternate
with them, as they normally should be. The same thing occurs
in Poppies and many other Papaveracez, also in some Cruciferee ;
and in some of the cases each stigma
is more or less two-lobed. ‘This sug-
gests the explanation,’ here probably
the true one, which supposes that the
placentze are borne on the leaf-margins
in the normal way, but that each
stigma is two-parted (as if the carpel-
lary leaf were deeply notched at the
apex, and so its two stigmatic leaf-
margins separate, as Drosera illus- Ss
trates, Fig. 553), and that the two oy Soa
half-stigmas of adjacent carpels have 338
coalesced into one body, which would
of course stand over the parietal placentze beneath. Each stigma
in such a case, as well as each parietal placenta, would consist
of the united margins of two adjacent carpels.

§ 2. IN GYMNOSPERMS.

504. GyMNosPERMOUS (that is, naked-seeded) plants are so
named because the ovules, or bodies which are to become seeds,
are fertilized by direct application of the pollen, which reaches
and acts upon the nucleus of the ovule itself, not through the
mediation of stigma and style. In the structure of their flowers,
these plants are of a low or simplified type, in some respects not
obviously homologous with the Angiosperms which now consti-
tute the immense majority of phanogamous plants. But, up to
a comparatively late geological period, Gymnosperms appear to
have been the only flower-bearing plants. They are represented

1 Given by Brown, in the Plante Javanice Rariores, above referred to.

FIG. 551. Transverse section of an ovary of Nymplza odorata, the carpels ovulifer-
ous over the whole interior surface.

FIG. 552. Pistil of Parnassix, with ovary transversely divided.

FIG. 553. Pistil of Drosera filiformis, with ovary transversely divided.

GYNCECIUM OF GYMNOSPERMS. 269

in the extant vegetable kingdom by three (or four) groups or
orders, two of them small, and one comparatively ample and of
wide distribution ; and all are so strikingly different from each
other that they cannot be illustrated by a common description.
The largest order, Conifers, is familiar, and contains a good
share of the most important forest trees of temperate climates.
The smallest, Gnetacez, chiefly tropical or of warm regions,
lies between Gymnosperms and common Dicotyledons. The
third, Cyeadaceze, is most remote from them, and as much so
from Monocotyledons, except that it imitates Palms, as it
also does the Tree-Ferns, in habit, both as to stem and _foli-
age. The particular morphology of Gymnosperms would re-
quire for its illustration copious details and the history of various
conflicting hypotheses. It must be relegated to the special
morphology of the natural orders, premising, however, a brief
sketch of the general floral structure.’

505. In Gnetaceer, Gymnosperms and Angiosperms almost
come together. The flowers have a perianth (diphyllous or
tetraphyllous) ; the stamens have a distinct filament and anther ;
and the gyneecium is a sac (presumably of two carpophylls)
open at the top and filled at bottom by a single oyule of the
simplest kind, 7. e. consisting of a nucleus destitute of coats.
This pistillary body is attenuated and prolonged above the ovule
into a style-shaped tube, with open and commonly two-cleft
orifice. In thealmost hermaphrodite sterile flower of Welwitschia,
this takes the form of a much dilated stigma, which is even beset
with seeming stigmatic papille. If only 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 morphoiogy still in part
unsettled, need not here be given. The history and the idea of gymnospermy
began with Robert Brown’s paper on Kingia, “ with Observations .. . . on
the Female Flower of Cycadeze and Conifer,’ 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. 435, and in Bliithendiagramme, i. 55-69; also ii. preface x.
See also Alph. DeCandolle, Prodr. xvi. 345, 524. In this volume, the late
Prof. Parlatore adhered to the ancient ideas in his monograph of the Conifere.

2 The view here implicitly adopted is that of Beccari, founded on the study
of Gnetum, and published in Nuovo Giornale 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. xxviii. 1872.

270 THE FLOWER.

genus Gnetum, shrubs or trees, with nearly the aspect of
Angiosperms, having broad and pinnately-veined leaves; Wel-
witschia of tropical W. Africa, remarkable for its persistent
cotyledons which form the only foliage of a woody and long-
enduring plant, and for its stem or trunk which broadens with-
out lengthening, except in its flower-stalks ; also Ephedra, of
much branched shrubs, mainly of warm-temperate regions, leafiess
or nearly so, one species of which inhabits Europe and two the
southern borders of the United States.

506. The flowers in all Gymnosperms are diclinous, either
dicecious or moneecious ; except that those of the strange Gneta-
ceous genus Welwitschia are structurally polygamous, the male
flowers having a well-formed but sterile gyncecium.

507. In Coniferx, the largest and most important type, are
embraced all the familiar Gymnosperms of temperate regions,
Pines, Firs, Cedars, Cypresses, which >
bear their flowers in catkin-like clusters {Y=
and their fruit in cones, and also the \¥,.
Yews and allied trees which do not/C-~
produce cones. Perianth being want-
ing and the sexes wholly separate, the floral type
is so degraded that it becomes doubtful whether
each cluster of anthers, or of ovuliferous seales
or ovules, constitutes a blossom or an inflores-
cence. Certain botanists look upon a _ whole
catkin, and others upon a male catkin only, of
a Pine or Fir as forming one flower. It is here
assumed that each stamen (

of the one and each oyu-
liferous scale of the other
answers to a flower of the
simplest sort.! The anthers
oS are extrorse, the cells or
pollen-sacs belonging to the outer or lower side of a scale or a

557

1 Tt will be seen that, for the female flowers, this follows of course from
generally accepted view ; and, where this is conceded, analogy may extend it
to the male catkins also: yet in such cases, where all the phylla of an indefiniie
simple axis are stamens, spirally arranged on it, the difference betwecn
inflorescence and male flower completely vanishes.

FIG. 554. Female flower of a Yew, an ovule surrounded by its bracts. 555. Longi-
tudinal and more enlarged section of a female flower of Yew and of the upper part of
the shoot it terminates: the thick coat of the ovule open at the top, the nucleus within,
and the beginning of the disk outside of the coat, are seen in section. After Stras-
burger.

FIG, 556. Young fruit (berry-like cup surrounding the seed) of Yew. 557. “Longi-
tudinal section of a mature fruit of the same. After Decaisne.

GYNGC!IUM OF GYMNOSPERMS. 271

connective: sometimes these sacs or cells are two, and the organ
evidently homologous with an ordinary stamen: often they are
more numerous (from e
three to twenty) and
yariously disposed.
508. The Yew Fam-
ily (Taxinez) is next to
Gnetacez in structure.
It is generally ranked as :
a suborder of Coniferze, 559
but it may claim to be a distinct order. The gynecium is a
naked ovule, terminating a stem,’ and surrounded by several
bracts. After fertilization, an outgrowth of the
receptacle (or a kind of disk, 394) makes its
appearance as a ring girding its base: this
grows in height and thickness, and becomes a
soft-fleshy cup, imitating a hollow berry, in
the bottom of which the stony-coated seed
nestles. (Fig. 554-557.) Very similar is the
gynecium of Torreya, except that the cup-
shaped disk develops almost simuitaneously
with the ovules, and as it grows becomes adnate
to the large seed in the form of a
fleshy coating. In the Gingko, two
or more similar ovules are nakedly
developed on a naked peduncle, un-
accompanied even by a bract (Fig.
598), and one or more of these ripens
into the berry-like seed, Fig. 559.
In Podocarpus there are some sub-
tending bracts, and the naked ovyule

1 Tt does not therefore follow that the ovule isa part of the axis, or is
terminal in the sense of being its direct continuation. In this regard it may
be only what the pistil of a Cherry is, which to all appearance is equally a
terminal production, but is really the representative of the last leaf of the
axis. If so, that leaf is here suppressed to the utmost, and replaced by
what is ordinarily its outgrowth, the ovular nucleus and its coat. The
structure of Podocarpus favors this interpretation.

FIG. 558. Female flowers of Gingko biloba or Salisburia adian ifolia. 5584. Portion
of the same enlarged. After Strasburger. 559. A drupaceous seed of the same, in
vertical section, exhibiting the mature disk which forms the flesh, the ernustaceous seed-
coat, within which is the keruel of the seed; at the base on one side a sterile ovule is
seen. After Decaisne.

FIG. 560. Female flower of Podocarpus (an oyule inverted on a column or elevated
support), subtended by bracts. After Eichler.

FIG. 561. Magnified vertical section of a similar flower of Podocarpus. After
Strasburger.

212 THE FLOWER.

is inverted on a more or less lengthened and stout support, which
is conceived to represent the carpel. (Fig. 560, 561.)

509. In the true Coniferze, to which Pines, Cypresses, and all
such cone-fruiting trees belong, the ovules are borne on or in
the axils of scales which are imbricated on a simple axis, in a
spicate or capitate manner; and the male flowers, each a single
stamen, are also similarly spicate or capitate. Both are com-
monly termed aments or catkins; and the female
ones properly so, according to the present view ;
but the only scales of the male catkins are parts
of the anther, being a dilated tip of the counective
in Pines, and a scale bearing anther-cells or pollen-
sacs on its back in Cypress.

510. In the Pine tribe the flowering female catkin consists of
bracts, spirally imbricated on the cauline axis: in the axil of
each bract or sterile scale is developed a scale
which bears two ovules, and is therefore regarded
as of carpellary nature. These ovules are pro-
duced on the lower part of the upper face of this
carpellary scale, and are wholly adherent to it
quite to the orifice, which is directed downward.
(Fig. 562, 563.) The ovuliferous scale in
becoming fructiferous usually much and soon out-
grows the bract, which is concealed in the Pine-
cone (or sometimes obliterated) ; but it remains
conspicuous in sundry Fir-cones. After fertil-
ization, the scales, successively covering each
other in close imbrication, protect the growing
seeds as effectually as would a closed ovary.
Sooner or later after ripening the scales diverge, and the seeds
peel off the face of the scale with a wing attached, and fall or
are dispersed by the wind.”

1 Among those who admit as well as those who reject gymnospermy,
there has been much controversy over the morphology of the parts. With
the former, the discussion turns on the character of the ovuliferous scale.
As to this, the hypothesis originally proposed by Mohl, and adopted by
Braun, is now said to be satisfactorily demonstrated by Stenzel, in Nov. Act.
Nat. Cur. xxxviii. 1876. See note by Engelmann in Amer. Jour. Sci. Dec.
1876, and also the preface to the second part of Eichler’s Bliithendiagramme,

FIG. 562. View of the upper face of a carpellary scale of a Larch, showing the pair
of adnate ovules.

FIG. 563. Similar view of a carpellary scale of a Larch, and of a bract behind if.
564. Ground plan of the same in diagram, reversed; the upper figure denoting the axis
of the cone, the lower the bract, the middle one the carpellary scale and the two ovules
borne on its face. After Eichler.

GYNCECIUM OF GYMNOSPERMS. 273

511. In the Araucaria tribe the ovuliferous or carpel-scale is
throughout smaller than the bract, and is completely adnate to
it, or with only the tip free; that of Araucaria (Fig. 565) bears
only one ovule, high on the carpel, the orifice downward as in
the Pine tribe. In Taxodium, Sequoia, and the like, the cone-
scale is equally inferred to be composed of bract and carpel-scale
united ; and indications of this composition are to be observed.
The ovules (from two to several) are at the base of the scale,
erect and free. The cone-scales are alternate and spiral on the
axis, but indistinctly so in Taxodium, the Bald Cypress or
so-called Cypress of the Southern United States.

512. In the true Cypress tribe (Cupressinez) the cone-scales,
which are never numerous, are opposite or verticillate, 7. e. like
the foliage-leaves, in whorls of twos, threes, or
sometimes fours; and the ovules are from two to

1878, where it is fully adopted. It was suggested by certain rather common
monstrosities, and by the two combined leaves of Sciadopitys.

According to this view, the ovuliferous scale in the Pine tribe is com-
posed of two leaves of an arrested and transformed branch from the axil of
the bract, which are in the normal manner transverse to the subtending
bract, are here carpellary, each bearing an ovule on the dors:/ face; the two
are coalescent into one by the union of their posterior edges, and the scale
thus formed is thus developed with dorsal face presented to the axis of the
cone, the ventral to the bract. It is therefore a compound open carpel,
composed of two carpophylls. This character of being fructiferous on the
back or lower side of the leaf occurs in no other phenogamous plants, but is
the rule in Ferns, from something like which Coniferee may be supposed to
have been derived; the ovules of the one in this regard corresponding to the
sporangia of the other.

FIG. 565. Vertical section (in diagram) of a bract, adnate carpel-scale, and adnate
ovule of Araucaria imbricata. After Eichler.

FIG. 566. Branchlet of the American Arbor-Vitz, considerably larger than in na-
ture, with a forming fertile cone. 567. One of the scales remoyed and more enlarged,
the inside exposed to view, showing a pair of naked erect ovules on its base.

FIG. 568. Fertile flowers of true Cypress (Cupressus sempervirens), after Baillon: a
forming cone, with one scale cut away, to show the cluster of ovules under it.

18

274 THE FLOWER.

several at or on the base of each cone-scale, always with orifice
upward. Arbor-Vite (Fig. 566, 567) has a single paix of ovules
to the scale; Junipers, sometimes only one; true Cypresses (as
in Fig. 568), often a dozen or more. At flowering time, the cone-
scales mostly appear as if simple ; but in most genera they soon
thicken greatly within; and they are usually understood to be
composed of bract and carpel-scale combined, the latter of the
same constitution as that of Pines and Spruces, but perfectly
consolidated and confluent with the bract-scale.?

513. In Cyeadacee, the type of the flower of Angiosperms is
almost or quite lost; yet the organs may be homologized with
those of Coniferze, which these plants are wholly unlike in habit.

1 This internal and ovuliferous scale may seem to be wholly hypetheti-
cal, and assumed to homologize the cupressineous with the abietineous
cone. Without it, we should have to consider that, while in Abietinee the
ovules belong to leaves of a secondary axis, in Cupressineew they are borne
on those of a primary axis, or else are axillary productions without carpels.
But in the Araucaria tribe the internal scale is obvious ; and there are sufii-

FIG. 569-575. Zamia, chiefly Z. media, after Richard. 569. A male plant. 570
Lower part of a male catkin. 571. A stamen removed, showing numerous small pollen-
sacs under the peltate top. 572. A female catkin, with a quarter section cut away.
573. A female flower or carpel, with two enlarging ovules or young seeds. 574. Ripe
seed, with the thick fleshy coat cut away at apex. 575. Longitudinal section of ripe
seed, more enlarged.

GYNGCIUM OF GYMNOSPERMS. 273

Their likeness to Palnis and other Monocotyledons is confined to
the port of their unbranched trunks and their pinnate leaves
with parallel-veined or simple-veined leaflets ; nor have they any
further resemblance to Ferns, except that in some the leaflets are
circinate in vernation. Although a tropical type (of small present
importance, compared with the part which it played in the
Devonian and Cretaceous periods), it has one small representa-
tive (Zamia media, the Coontie) at the south-eastern extremity of

576 577 578

the United States, and a more striking one (Cycas revoluta, well
known in cultivation) in the southern parts of Japan.

514. Following the analogy of Conifer, each scale (whether
of the pollen-bearing or the oyule- 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, are exter-

cient indications of similar composition in the cupressineous cone-scales to
induce the adoption of it by Parlatore, who rejected the idea of gymnospermy ;
and, finally, this composition is nearly demonstrated by Van Tieghem (1868)
upon the anatomical structure, and by Strassburger (1872) on the development.

FIG. 576-578. Carpophylla of Cycas revoluta, much reduced in size. 576. One
bearing ovules below and leaflets or leaf-lobes towards the apex. 577. A similar carpo-
phyll with leaf-lobes reduced to mere teeth, and ovules in place of the lower teeth.
578. A similar carpophyll in mature fructification, bearing the large drupaceous naked
seeds. The last two after Richard.

276 THE FLOWER.

nally much alike in the two sexes, which throughout the family
occupy separate plants. The male flower (Fig. 568) or stamen,
if it may be so termed, bears indefinite pollen-sacs on the under
side of the peltate portion, sometimes extending to che upper
part of its stalk. The homologous female flower, or carpophyll,
bears a suspended ovule on each side of the stalk (Fig. 573),
which becomes a large flesny-coated seed. In Cycas the male
ament is not very dissimilar, although on a larger scale. But
the carpophylls are evident leaves, not condensed into an ament,
but loose or spreading, of a character and aspect intermediate
between the lax bud-scales which precede and the pinnate foliage-
leaves which follow them in development. Along the margin
of what would be leaf-blade they bear ovules in place of leaflets,
lobes, or teeth (Fig. 576-578) ; and these, when fertilized from
the male flowers, mature into large and drupaceous naked seeds.
Even without fertilization, such seeds grow to their full size on
the female plant of the common Cycas (or falsely so-called Sago
Palm), but form no embryo.

Section VIII. Tue Ovoute.!

515. Ovules (302) are peculiar outgrowths or productions of
earpels which, upon the formation of an embryo within, become
seeds. In the angiospermous gynecium (476) they are nor-
mally produced along the margins, or some part of the margins,
of the carpellary leaf (478), either immediately, or by the in-
termediation of a placenta (485), which is a more or less evident
development of the leaf-margins for the support of the ovules.
Rarely, yet in a considerable number of cases (501, 502), ovules
are developed from the whole internal surface of the ovary, or
from various parts of it, in no definite order, directly from the
walls, and without the intervention of any thing which can be
regarded as placenta. In Gymnosperms (504-514) the ovules
are borne on the face of the carpellary scale or at its base; or
on leaf-margins, as in Cycas; or, when there is no representa-
tive of the carpel, on the cauline axis, seemingly as a direct
growth of it. (508, note.)

516. As to attachment. ovules are either sessile, 7. e. stalk-
less, or on a stalk of their own (Fig. 582, 584), the FuNicuLUs
or Poposperm. As to number they are either solitary, few, or
en ee Se ee

1 Lat. Ovulum, pl. Ocula, diminutive of ovum (egg), perhaps first used by
Adanson

OVULES. ZT

indefinitely numerous. They may also be indefinite or variable
in number when not particularly numerous.

517. As to situation and direction within the ovary, the terms
are somewhat special. Ovules are ereef, when they rise from
the very bottom of the cell, as
in Fig. 580; ascending, when
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
Nuctevus. This in most cases is invested by one or two proper
coats. The coats are sacs with a narrow orifice, the ForaAMEN.
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 Exosrome, of the inner ENDOSTOME ;
literally the outer and the inner
orifice. The coats themselves have
been named Primrye and SrEcun-
DINE, but with an ambiguity in the
application which renders these
names unadvisable: for in their »—..
formation the coats appear later
than 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 Cuataza. The attachment of the
ovule to its funiculus or support, which in the seed becomes the

379

J Hs

582 oS4

FIG. 579. Ovary of a Buttercup, divided lengthwise, to display its ascending ovule.
580. Same of Buckwheat, with an erect ovule. 581. Same of Anemone, with a sus-
pended ovule.

FIG. 582. Diagrammatic section of a typical or orthotropous ovule (such as that of
Fig. 5824), showing the outer coat, a, the inner, b, the nucleus, c, the chalaza, or place
of junction of these parts, d. (The coats are never so separated and the nucleus so re-
duced in size as is represented in this mere diagram.) 583. An ovule similar to the
preceding, but curved, or campylotropous. 584. An amphitropous oyule.

* 278 THE FLOWER.

HixvM, takes also this latter name in the ovule. In the simplest
form of ovule (as in Fig. 582, 580), hilum and chalaza are one.
So also in cases where the body of the ovule incurves, as in Fig.
583. But very commonly the place of attachment, which becomes
the hilum, is more or less distant from the chalaza ; as in Fig. 584
and 587, where the hilum is lateral, but the chalaza at the larger
end, the two being connected by a short ridge; and in Pig. 588
the two are separated by the whole length of the ovule.

519. The simplest and most rudimentary oyule is that with-
out a coat, as in Mistletoe and the whole order Loranthaceze, and
in Santalacezee and Gnetacee. This has been called a naked
ovule; but long before ovules of such simplicity were known
this term had been appropriated to those of Gymnosperms, in
the sense of destitute of ovarial or pericarpial covering, 7. e. to
uncovered ovule, not to uncovered nucleus. The oyule consist-
ing only of nucleus may be termed (after Alph. DeCandolle)
simple, or better achlamydeous.*

520. The tunicated or chlamydeous oyule is of three principal
kinds, with one or two subordinate modifications. These are
the orthotropous, campylotropous, and anatropous, and the modi-
fication called half-anatropous, or amphitropous.”

521. Orthotropous (Fig. 580, 582, 585), or straight ovule, is
the simplest but least common species, being that in which the

chalaza is at the evident base, and the orifice at the opposite
extremity, the whole ovule straight (as the first part of the name
denotes) and symmetrical. Atropous, meaning not turned at
all, is a later and etymologically much better name, but it has

1 An epidermal stratum or tegument may not be wanting to such ovules,
forming a sort of adherent covering; but this in nature and origin is not
similar to the ovular coats.

2 In Latin form, orthotropa, campylotropa, anatropa, amphitropa, — names given’
by Mirbel, and referring to the way in which the ovule is turned either on
itself or on its support. Some English botanists incongruously write ortho.
tropal, campylotropal, &c.

FIG. 585. Orthotropous or Atropous ovule of Buckwheat. 586. Campylotropous
ovule of Chickweed. 587. Amphitropous ovule of Mallow. 588. Anatropous ovule of a
Violet. The letter k indicates the hilum; c, the chalaza, which in 585 and 586 corre-
sponds to the hilum; /, the foramen or orifice; 7, the rhaphe.

OVULES. 279

not come into general use. This ovule is characteristic of
Polygonacez, the proper Urticaceae, Cistacer, &e.

522. Campylotropous (Fig. 585, 586) is the name of the ovule
which in the course of its growth is curved on itself so as to
bring the orifice or true apex down close to the base, here both
chalaza and hilum. ‘This and the orthotropous ovule begin
their development on the placenta in the same way, but the
campylotropous develops unequally, one side enlarging much
more than the other, especially at the base, until the ovule
becomes reniform, and chalaza and orifice are brought into
close proximity. Campylotropous ovules are characteristic of
Cruciferze, Capparidaceze, Resedaceze, Caryophyllacez, and
Chenopodiacez.

523. Amphitropous (Fig. 584, 587), also termed Heterotropous
and sometimes Half-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 very 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 Primulacez, and
is common in Legumminos.

524. Anatropous (Fig. 588, also 579, 581, 597) is the name
of far the commonest species of ovule, that in which the organ,
under the course of its growth, is quite inverted on its base ; so
that, instead of standing at right angles with the funiculus, it is
parallel with it, or rather with the apparent continuation of
it, which is adherent to its surface as a sort of ridge or cord
extending along the whole length of the ovule, from hilum to
chalaza. The latter occupies the seeming apex of the seed ;
and the organic 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 cord on that side not rarely prominent.

525. The cord or ridge, which extends along the whole length
of the anatropous ovule, and for half its length in the amphi-
tropous (Fig. 588, 587, 7), is named the Ruapure. This is not

280 THE FLOWER.

at all a seam, as the Greek word denotes. Its origin, and the
whole structure of such ovules will be apprehended by comparing
various stages of its growth.

526. An ovule of any kind at the beginning is an
excrescence or outgrowth of the placenta, or of some
part of the leaf-surface if there is no developed
placenta. This incipient oyule is the nucleus (518),
or the nucleus surmounting a rudimentary funiculus,
The nacleus is soft cellular tissue only, from first to
last. The achlamydeous ovule (519) undergoes no
further development except in size or shape. Indeed
sometimes (as in Balanophoreze) this bare nucleus
is reduced to a few cells of parenchyma.

527. In ordinary ovules a new growth early begins around
the base of the nucleus, or is sometimes coetaneous with it, at
first as a ring (or part of a ring), soon as a cup, at length as
an enclosing sac or covering, open at the top; this is the inner
coat of the ovule when there are two. ‘The outer coat begins and
goes on in the same way, and at length grows over and encloses
the inner coat as that did the nucleus. (Fig. 590-595.) When-
ever there is a third and more exterior coat it is formed during
the growth of the fertilized ovule into the seed, to which there-
fore it belongs, and in which it takes the name of arillus. (597.)
At the time of fertilization the apex of the nucleus, or a pro-
longation of it, usually projects beyond the orifice and there
receives the descending pollen-tube. Some fibro-vascular tissue,
especially spiral ducts, may be found in the funiculus and cha-
laza, sometimes extending into the coats.

528. The development of the orthotropous or atropous (un-
turned) ovule proceeds symmetrically, without distortion, the

parts keeping their primitive direction. In the campy lotropous,
the whole of one side of the ovule greatly outgrows the other.

FIG. 589. Magnified view of a vertical section of a carpel of Magnolia Umbrella,
about a month before anthesis, showing one of the two nascent ovules, at this time
only nucleus.

FIG. 590-597. Further development of the ovule of Magnolia Umbrella, showing the
formation of the coats and the anatropy. 590. Ovule a week older than in 589. 591.
Same a week or two later. 592. Same a few days later 593 Same from a nearly full-
grown flower-bud. 594. Same at time of anthesis. 595. Vertical section of the last through
the middle of the rhaphe. 596. Cross-section of the same. (See Jour. Linn, Soc. ii. 108:)

OVULES. 281

In the anatropous, the inequality of growth is mainly confined
to the base or chalazal region, which ends by becoming upper-
most; and the full-grown oyule has the ap-
pearance of being inverted on and. adherent
to the upper portion of its funiculus, the
rhaphe. Fig. 589-597 illustrate the course
of development from a comparatively early
period.

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 597
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 are solitary or in single rows, and
either ascending or hanging, the rhaphe is usually on the side
next to the placenta or ventral suture, as in Fig. 579: it is then
said to be ventral (7. e., next the ventral suture), or adverse to
the placenta. In certain cases, mostly in hanging ovules, as in
Fig. 581, 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.+

1 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. 581. Upon this
conception, Euphorbia and its allies has normally suspended ovules, the
rhaphe being next the placental axis, and Buxus and its allies, resupinately
suspended ovules, the rhaphe averse. The propriety cf 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 Rhamnus and its allies; and here it was shown
by Bennett (in Pl. Javan. Rar. 151), and confirmed by the analyses of
Sprague (Gray, Gen. III. ii. 168, plates 163-169), 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. Pl. 178, &c.) maintains the
contrary, but 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. 597. Same as 595 more magnified; the outer coat (a). the inner (b), nucleus (ce),

and the bundle of spiral ducts (() in the rhaphe (running from placenta to chalaza)
indicated.

282 THE FLOWER.

530. Origin and Nature of the Ovule. It has been already
stated in general terms that ovules are peculiar outgrowths
or productions, generally of the margins of carpellary leaves
(515) ; that they are composed of parenchymatous cellular sub-
stance, at least as to the nucleus, of which the simplest ovule
wholly consists (526) ; that the coats originate subsequently to
the nucleus ; and that the outer coat is of later origin than the
inner one. (518.) The mamiliform protuberance of which the
forming ovule at first consists originates in one or more cells of
a layer directly beneath the epidermis.*

531. The morphological nature of the ovule has been much
discussed. ‘The commonly prevalent view was that the ovule
is homologous with a leaf-bud, and that its nature is in some
degree illustrated by such buds as those which develop on the
margins of the leaves of Bryophyllum, as shown in Fig. 322.
But such buds, and the bulblets or fleshy buds which appear on
the face of certain leaves, follow the universal order of budding
growth, that is, are centripetal in development, the outermost
parts being the earlier and the inmost the later formed. The
ovule, on the contrary, is basipetal or centrifugal in develop-
ment, the nucleus being first and the outer coat last formed ;
therefore the coats are not homologous with sheathing leayes,
nor the nucleus with a vegetative axis. The older theory has
accordingly given way to the present one, in which the ovule
answers to the lobe of a leaf peculiarly transformed, or to an
outgrowth of a leaf, whether from its edges or surface. The

apotropous, epitropous, and heterotropous (the first two new, the last employed
in a new sense), the more so since the application is confused with hypo-
thetical considerations and the necessity of bringing the ovules ideally back
to ascending or horizontal positions. It may be stated, briefly, that /Zetero-
tropous, in Agardh’s terminology, applies to the normal position of collat-
eral ovules, with rhaphes back to back, in opposite directions on the two
halves of the placenta; Apotropous, to an erect or ascending ovule with its
rhaphe next the placental axis, and a hanging one has its rhaphe averse
from it; /pitropous, when an erect or ascending ovule has its rhaphe averse,
and a hanging one has it adverse.

1 Hofmeister’s statement that the simple ovule of Orchis originates in
the division of a single epidermal cell (and is therefore a trichome) is con-
troverted by Strasburger and by Warming. The latter adds the remark,
that even if it were so in cases of extreme simplicity, this would not invali-
date the proposition that the ovule is to be regarded as the homologue of the
lobe of a leaf. Such a lobe is not rarely reduced to a single bristle. For
the whole subject of the origin, development, morphology, and theory of the
ovule, see Warming’s very elaborate and perspicuous memoir, De l’Ovule;
also the papers of Celakowsky, Van Tieghem, &c., referred to in notes to
paragraphs 500, 501.

OVULES. 283

ereat advantage of this view is that it serves to homologize the
fructification of Flowering Plants with that of the higher klower-
less Plants, or the Ferns, the sporangia or analogues of the
ovule being outgrowths of the leaf.!

532. Origination of the Embryo. ‘The whole process of fer-
tilization and the resulting produc- ee
tion of the embryo, also the history Y a
of the subject, belongs to the suc-
ceeding volume, involving as they
do questions of minute anatomy and
of physiology. But a general idea
may here be given of the way in
which the embryo originates. The
tube which a grain of pollen sends
forth into the stigma (074, 575)
penetrates the style through loose
conducting tissue charged with
nourishing liquid, reaches the cavity
of the ovary, 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 embryo-
sac, which is formed by the great
enlargement of a single cell of the
tissue, or of two or more cells the ,
product of a mother cell) forms in
the nucleus, the upper part of it
commonly reaching nearly or quite
to the apex of the nucleus, which
the pollen-tube impinges on or
sometimes penetrates. A particular portion of the protoplasm
contained in the embryo-sac forms a globule, and this at the time

! The advocates of this view naturally maintain that ovules and placente
always belong to leaves, and never truly to a cauline axis; that in the pre-
central placentation of Primulacez, the actual ovuliferous surface is an out-
growth of the bases of the carpellary leaves coalescent with each other and
adnate to a prolongation of the torus ; also that in those Gymnosperms which
have no carpophyll, such as Yew, the whole nascent carpellary leaf, or rather
the papilla which would otherwise develop as such, is directly developed into
ovule. This, being solitary and the last production of the axis, necessarily
appears to terminate it. (500, 501, notes.)

FIG. 598. Diagram representing a magnified pistil of Buckwheat, with longitudinal
section through the axis of the ovary and orthotropous ovule; some pollen on the stigmas,
one grain distinctly showing its tube, which has penetrated the style. reappeared in the
cavity of the ovary, entered the mouth of the solitary ovule (0), and reached the
embryo-sac (s) near the embryonal vesicle (7).

284 THE FLOWER.

of fertilization is found at the apex of the sac, at or adjacent to
the part reached by the pollen-tube. Not rarely it adheres to the
599 600 601 wall of the sac exactly
opposite the termination
of the pollen-tube. This
is called the embryonal
vesicle. To it the con-
tents of the pollen-tube
are in some manner trans-
ferred. Upon which it
takes a more definite
shape, acquires a wall of
cellulose, and so becomes
a vegetable cell. This
divides into two, the lower
again into two, and so on, forming a chain (the swspensor or pro-
embryo). The terminal cell of this divides again and again in
three directions, producing a mass of cells which shapes itself
eos 605 606 into the embryo, the initial
plant of a new genera-
tion. Ordinarily the sus-
pensor soon disappears.
It is attached to the ra-
dicular end of the em-
bryo, which consequently always points to the foramen or
micropyle of the seed. The process in Gymnosperms is more
complex, and has to be separately described.

533. Polyembryony, the production of two or more embryos in
one seed, is not uncommon in Gymnosperms (there being a kind
of provision for it), and is of occasional but abnormal occurrence
in Angiosperms, in the seed of Mistletoe, Santalum, &c. In
these it results from the production and fertilization of more
than one embryonal vesicle. Strasburger has recently ascer-
tained that the commoner polyembryony in the seeds of Onions,
Oranges, Funkia, &c., results from the production of adventive
embryos, which originate in the nucleus outside of the embryo-
sac and wholly independent of fertilization.1 Two kinds of

1 Strasburger, Ueber Polyembryonie, in Zeitschr. Naturwis. Jena, xii.
1878 (see Amer. Jour. Sci. April, 1879). It was found that when, by exclu-
sion of pollen, the formation of a normal embryo was prevented, no adventive

FIG. 599. Diagram of the suspensor and incipient embryo at its extremity. 600.
The same, with the embryo a little more developed. 601. The same, more developed
still, the cotyledons faintly indicated at the lower end. 602. Same, with the incipient
cotyledons more manifest. 603. The embryo nearly completed.

FIG. 604-606. Formingembryo froma half-grown seed of Buckwheat, in three stages.
607. Same, with the cotyledons fully developed.

THE FRUIT. 285

anomalous reproduction are therefore now known, which are
intermediate between sexual and non-sexual, between budding
and fruiting propagation, viz., —

Apogamy, which is budding growth or prolification in place of
that which should subserve sexual reproduction. This was dis-
covered in Ferns by Prof. Farlow, while a pupil of De Bary, by
whom our knowledge of the process has recently been extended,
and this name imposed.’ The production of bulblets in place of
seed or embryo answers to this in Flowering plants.

Parthenogeny, the counterpart analogue of apogamy, is the
non-sexual origination of an embryo extraneous to the embryonal
vesicle or even the embryo-sac. However abnormal, its occur-
rence is probably not so rare as has been supposed.

GHAPTER VIL.
THE FRUIT.

Section I. Irs Structure, TRANSFORMATIONS, AND DEHISCENCE.

534. The Fruit consists of the matured pistil or gyncecium
(as the case may be), including also whatsoever may be joined
to it. It is a somewhat loose and multifarious term, applicable
alike to a matured ovary, to a cluster of such ovaries, at least
when somewhat coherent, to a ripened ovary with calyx and
other floral parts adnate to it, and even to a ripened inflores-
cence when the parts are consolidated or compacted. Fruits,
accordingly, are of various degrees of simplicity or complexity,
and should be first studied in the simpler forms, namely, those
which have resulted from a single pistil. Such a fruit consists
of Pericarp with whatever may be contained in it and incorpo-
rated with it.

embryo appeared in those seeds which habitually produce them. To this
Celebogyne offers an exception. The female of this dicecious plant habit-
ually matures fertile seeds, with a well-formed embryo, in Europe when there
are no male plants in the country. Strasburger ascertained that the embryo
thus formed is adventive, the embryonal vesicle perishing. Purthenogenesis,
of which Czlebogyne was the most unequivocal] case, is thus confirmed, and
is shown to occur in most polyembryony ; but it is at the same time explained
to be a kind of prolification.

1 See Farlow, in Proc. Am. Acad. ix.68; De Bary, Bot. Zeit. xxxvi. 465-487.

286 THE FRUIT.

535. The Pericarp, or Seed-vessel, is the ripened ovary. It
should, therefore, accord in structure with the ovary from which
it is derived. Yet alterations sometimes take place during fruc-
tification, either by the abortion or obliteration of parts, or by
accessory growth.

536. Internal Alterations. Thus, the ovary of the Oak con-
sists of three cells, with a pair of ovules in each; but the fruit
has a single cell, filled with a solitary seed, only one ovule being
matured, while two cells and five ovules are suppressed, the
remains of which may be detected in the acorn. The ovary
of the Chestnut has six or seven cells, and a pair of suspended
ovules in each; but only one of the dozen or fourteen ovules
ever develops into a seed, except as a rare monstrosity. The
three-celled ovary of the Horsechestnut and Buckeye is similar
in structure (Fig. 608-611), and seldom
ripens more than one or two seeds; but the

abortive seeds and cells are obvious in the ripe fruit. The
ovary of the Birch and of the Elm is two-celled, with a single
ovule in each cell: the fruit is one-celled, with a solitary seed ;
one of the ovules being uniformly abortive, while the other in
enlarging thrusts the dissepiment to one side, and obliterates the
empty cell. Similar suppressions in the fruit of parts actually
extant in the ovary are not uncommon.

537. On the other hand, there may be more cells in the fruit
than there are primarily in the ovary. Thus the fruit of Datura
is dicarpellary and normally two-celled, with a large placenta
projecting from the axis far into the cells. But each cell be-
comes bilocellate, that is, divided into two, by a false partition
growing out from the back of each carpel and cohering with the
middle of the adjacent placenta. So the 5-carpellary and nor-
mally five-celled ovary of common Flax early becomes spuriously
ten-celled (morphologically speaking, not 10-locular, but 10-
locellate), by a false partition extending from the back of each

FIG. 608. Longitudinal section of the ovary of a Buckeye (sculus Pavia), showing
the pairs of ovules in two of the cells. 609. Transverse section of the same displaying all
three cells and six ovules. 610. Same of half-grown fruit, with single fertile seed, abor-
tive ovules and obliterating cells. 611. Dehiscent one-seeded fruit, diminished in size.

ITS STRUCTURE AND TRANSFORMATIONS. 287

carpel across its cell (Fig. 539-541) ; and the solitary carpel
is similarly divided lengthwise in many species of Astragalus,
as in Fig. 534. Transverse divisions or constrictions across
a maturing ovary (such as is seen in Fig. 620) are not uncom-
mon, especially in legumes and other pods, and are of little mor-
phological significance.

538. External Accessions may here be referred to. The wing
of the pericarp in Maple, Ash, and the like (Fig. 625-627), are
familiar instances of this; and of the same nature are the im-
bricated scales which cover some Palm-fruits; the prickles on
the pod of ‘Datura, Ricinus, &c., and the hooked or barbed
prickles of many small pericarps (as in various Borraginacez) ,
which thus become burs and are disseminated by adhering to the
hairy coat of cattle. All these are of the nature of superficial
outgrowths, and these especially affect the pericarp or parts
connected with it.

539. Persistence of Connected Organs. An adnate calyx (331),
being consolidated with the ovary, necessarily makes a constit-
uent part of the fruit, in the pome (575) doubtless a very large
part. The limb or lobes of such adnate organ may persist, as
the tips of the sepals on an apple or quince, and may be turned
to useful account, as is the pappus of Compositz for dissemina-
tion. Or, in small pericarps, the style may persist as part of the
fruit, and subserve the same ends, either by becoming feathery
for aerial dissemination, as in Clematis and in one section of
Geum, or by becoming hooked at the tip for adhesion to fleece,
&c., as in other species of the latter genus. Or adjacent parts
which are not actually incorporated with the pericarp may play
similar parts in the economy, as the hooks on the calyx-tube of
the dry calyx of Agrimonia, which at maturity is detached with
the included frait, the fleshy fructiferous calyx of Gaultheria (Fig.
651) and of Mulberry (Fig. 654); and the pulpy fructiferous re-
ceptacle of the strawberry (Fig. 653): the ultimate utilities in
both classes of instances being similar, viz., wide dispersion of
the seed by animals, whether by external carriage, or by being
devoured and the voided seeds of fleshy fruits thus disseminated.

540. Transformations in Consistence. In the change from
ovary to mature pericarp, various kinds of transformations may
take place. In some the wall of the ovary remains thin and
becomes in fruit foliaceous or leaf-like, as in a pea-pod, the
earpels of Columbine, and Marsh Marigold (Caltha), or the pod
of Colutea or Bladder Senna. In others it thickens and becomes
at maturity either dry throughout, as in nuts and capsules; or
fleshy or pulpy throughout, as in berries; or hard-rinded with-

288 THE FRUIT.

out but soft within, as in a pepo; or fleshy or berry-like without,
but indurated within, as in all stone-fruits, such as the cherry
and peach.

541. When the walls of a pericarp consist of two layers of dis-
similar texture (as in a peach) the outer layer is called Exocarp,
the inner Enpocarp, these terms meaning exterior and interior
parts of a fruit. When the external layer is a comparatively thin
stratum or film, it is sometimes termed the Ericarp. When it is
fleshy or pulpy it is named Sarcocarr. When the endocarp within
a sarcocarp is hard and bony or crustaceous, forming a shell or
stone, thisis termed a PurameN. When three concentric layers are
distinguishable in a pericarp, the middle one is called Mresocarp.

042. Fruits may be divided into two kinds, in reference to
their discharging or retaining the contained seeds. They are
dehiscent when they open regularly to this end; cndehiscent when
they remain closed. There is a somewhat intermediate condi-
tion, when they rupture or burst irregularly, as in Datura Metel,
&c. Dry pericarps with single seeds are commonly indehiscent ;
those with several or many seeds mostly dehiscent. Seeds pro-
vided with a wing or coma or any analogous help to dispersion
are always in indehiscent pericarps. Permanently fleshy peri-
carps are indehiscent, stone-fruits as well as berries. But in
some stone-fruits (7. e., with indurated endocarp and fleshy
exocarp), such as those of Almond (Fig. 640) and Hickory,
the barely fleshy exocarp or sarcocarp dries or hardens, instead
of softening, as maturity is approached, and at length separates
from the putamen by dehiscence.

543. Dehiscence, the opening of a pericarp for the discharge
of the contained seeds, is ~-gular or irregular; or, better, is
normal and abnormal. 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 in
general is that of Pop.

544. Regular or normal dehiscence is that in which a pericarp
splits vertically, for its whole or a part of its length, on lines
which answer to sutures or junctions, that is, along lines which
correspond to the margins or midribs of carpellary leaves, or to
the lines and surfaces (or commissures) of coalescence of con-
tiguous carpels. The pieces into which a pericarp is thus sun-
dered are termed VALves.

545. The normal dehiscence of a carpel is by its immer, ven-
tral, or ovuliferous suture, that is, by the disjunction of the
leaf-margins, as in Fig. 618. Its only other line of normal
dehiscence is by the opposite or dorsal suture, that is, down

DEHISCENCE. 289

the midrib. Legumes usually dehisce by both sutures (as in
Fig. 619), therefore into two valves.

546. A dehiscent pericarp formed of two or more carpels is
called a CapsuLe. The two leading terms descriptive of capsular
dehiscence were based upon the modes of opening of pericarps
having as many cells as carpels: they are the septierda/, that is,
as the term denotes, cutting through the septa or dissepiments ;
and the loculicidal, that is, cutting into the /oculi or cells.

547. Septicidal, the dehiscence through the dissepiments, is
the disjunction of a pericarp into its constituent carpels, these

then usually themselves dehiscing down their ventral suture,
as in Fig. 612, illustrated by
the diagram, Fig. 613. Good
‘examples are furnished by the
Hypericum Family (the pistil
illustrated in Fig. 536, 537),
where the placentae which
compose the axis are carried
away on the edges of the par-
titions or introflexed valves ;
also by Rhododendron, Kal-
mia, and the like, in which
the placentee remain combined.
into a column in the axis (the
CoLuMELLA or column), from
which the edges of the valves
break away.

548. The septicidal dis-
junction of the carpels does {| © Z
not of itself open the cells.
Such separated carpels when

aa one-seeded not rarely remain “_|_~
closed, as in Mallow, Ver- op
bena, &e. Or when dehiscent they may open both by the ventral
and dorsal sutures; 7. e., the pericarp may first. divide into its
constituent carpels, and then each carpel break up into half
carpels, as in Euphorbia.

549. Loculicidal, the dehiscence into the loculaments, loculi,
or cells of the pericarp (shown in Fig. 614, and the diagram,
615), is that in which each component carpel splits down its

FIG. 612. Septicidally dehiscent tricarpellary capsule of Elodes Virginica. 613. Dia-
gram of septicidal dehiscence.

FIG. 614. Loculicidally dehiscent tricarpellary capsule of an Iris, divided trans-
versely at the middle. 615. Diagram of loculicidal dehiscence.

290 THE FRUIT.

dorsal suture, as in Iris, Hibiscus, G£nothera, &c. In this, the
dissepiments remain intact. If they break away from the centre
then they are borne on the middle of the valves, as in the figures
above cited. If they remain coherent in the axis but break away
from the valves, the result is one form of what is called —

550. Septifragal dehiscence, ¢.e., a breaking away of the
valves from the septa or partitions, as shown in Fig. 616. This
represents the loculicidal form of the septifragal mode, which is
less common than that of the accompanying diagram, Fig. 617.

Here the partitions alternate

Pa YN __ With the valves; that is, the’

dehiscence of the pericarp is

of the septicidal order, as

near as may be, but the par-

titions do not split, wherefore

the valves break away at the

ae mm common junction. To this

the term marginicidal has been applied. It occurs in the 2-3-

carpellary capsule of Ipomeea (especially in the common Morning

Glory), in the 5-carpellary capsule of the North American species

of Bergia; likewise in the 2-carpellary pod of Crucifere (Fig.

623), with a difference that the placentz from which the valves
break away are here parietal and the partition is abnormal.

551. The terms septicidal and loculicidal apply equally in plan,
though not with etymological correctness, to one-celled capsules
with either parietal (495) or free central (599) placentee. When
the dehiscence is of the septicidal type and the placentation pari-
etal, the (half) placentze are borne on the margin of the valves,
as in the Gentian family and the species of Hypericum with one-
celled capsule. When the placentz are borne on the middle
of the valves, as in Violets, the dehiscence is of the loculicidal
type. In the case of free central placentze with no trace of
partitions, the character of the dehiscence may usually be deter-
mined by the position of the styles or stigmas relative to the
valves.

552. Dehiscence may be quite normal although very partial,
as when confined to the apex of the capsule of Cerastium and
of Primula, and even to the pores under the radiate stigmas
of Poppy.

553. Irregular or abnormal dehiscence is such as has no respect
to the normal sutures; as where the dehiscence is transverse ;

FIG. 616. Diagram of loculicidally septifragal dehiscence. 617. Same of septicidally
or rather marginicidally septifragal dehiscence.

a

ITS KINDS. 291

either extending part way round, as in the pod of Jefiersonia,
or completely round, so that the upper part falls off like an
unhinged lid. This ezrewmscissile dehiscence occurs in many
plants of widely different orders ; such, for example, as Purslane
(Fig. 621), genuine Amaranths, Plantain, Pimpernel, and Hen-
bane. In other cases, as in Antirrhinum (Snap-dragon) and
its allies, the cells burst by irregular laceration at a definite
point, and discharge the seeds through the ragged perforation ;
or one or more neat valvular orifices are formed on some parts
of the wall, as in Campanula.

Section II. Tue Krinps or Frutr.

554. Fruirs have been minutely classified and named ;! but
the terms in ordinary use are not very numerous. <A rigorously
exact and particular classification, discriminating between the
fruits derived from simple and from compound pistils, or between
those with and without an adnate calyx, is too recondite and
technical, and sometimes too hypothetical, for practical pur-
poses. It is neither convenient nor philosophical to give a
substantive name to every modification of the same organ. For
all ordinary purposes, both of morphological and systematic
botany, it will suffice to characterize the principal kinds under
the four classes of —

Simple fruits, those which result from the ripening of a single
pistil ;

Aggregate, those of a cluster of carpels of one flower crowded
into a mass;

Accessory or Anthocarpus, where the principal mass consists
of the surroundings or support of either a simple or an aggregate
fruit ;

Multiple or Collective, formed by the union or compact aggre-
gation of the pistils of several flowers, or of more than one,

555. Simple Fruits may be distinguished, upon differences of
texture, into Dry Fruits, Stone Fruits, and Baccate Fruits; or,
better, into Dry and Fileshy ; and the first may be divided into

1 The greater part of the forty-three substantive names of Desvaux’s,
and even of the thirty-six of Dumortier’s and of Lindley’s elaborate classi-
fications of fruits have never found employment in systematic botany, and
doubtless never will be used. Yet a detailed carpological classification has
its uses for the student. Among the more recent attempts are the successive
ones of Dickson, McNab, and Masters. See Nature, iv. 347 (also in Trimen’s
Jour. Bot. 1871, 310), iv. 475, and v. 6.

292 THE FRUIE.

dehiscent and indehiscent kinds.! Theoretically, each kind may
be divided into those of a simple and those of a compound pistil,
and some would make the primary division on this character.
Some also would separate fruits with adnate or superior calyx
from those free of all such combination. But in practice these
differences can seldom be indicated by substantative names.
The name of berry is equally applicable to the fruit resulting
from the single carpel of Actza, the syncarpous ovary of the
grape, and the similar ovary with adnate calyx of a gooseberry
and cranberry. It should be, understood that the kinds shade
off one into another most freely.

556. Dehiscent Fruits (543), or Pods, are distinguishable into
apocarpous, or of single carpels, and syncarpous, of more than
one earpel, 7. e. the first of a simple, the second of a compound
pistil. The first kind is mainly represented by the olliele and
the Legume; the second, by the Capsule and its modifications.

557. A Follicle is a pod formed
of a simple pistil, and dehiscent by
one suture (this almost always
the ventral or inner suture) alone ;
as in the Larkspur, Columbine,
Peony, and Marsh-Marigold (Fig.

sa 618) ; also in Milkweed and Dog-
bane. There may be several follicles or only
one to a flower, even in the same genus, as in
Larkspurs, Cimicifuga, &c. In Magnolia
(Fig. 648-650), fleshy carpels become follicles
dehiscent by the dorsal suture.

558. A Legume is the pod formed of a
simple pistil which is dehiscent by both sut-
ures (as in the Pea, Fig. 619), so dividing
into two pieces or valves. (544.) This is the
fruit of the Pulse Family, accordingly named
Leguminosze (Leguminous plants): indeed,
the name of legume is restricted to the fruits of this family,
and in descriptive botany is extended to all the modifications

1 Dr. Masters’s modification of Dickson’s and McNab’s classification of
simple fruits, as to primary kinds, is into

1. Nuts, or Achcenocarps, dry and indehiscent ;

2. Pods, or Reqmacarps, dry, dehiscent ;

3. Stone-fruits, or Pyrenocarps, fleshy without, indurated within, indehiscent ;

4. Berries, or Sarcocarps, fleshy throughout, indehiscent.

FIG. 618. A dehiscent follicle of Marsh-Marigold, Caltha palustris.
FIG. 619. Legume cf a Sweet Pea, already dehiscent. 620. Loment of a Desmodium,

ITS KINDS. 293

which that order presents. Some of these, in fact, are in-
dehiscent and reduced to akenes; some break up at maturity
into one-seeded indehiscent articulations or joints, which are
dispersed as if they were so many seeds. A legume of the latter
kind takes the special name of Lomenr, Lat. Lomentum. (Fig.
620.) In Mimosa (Sensitive-plant, &c.), such articulations de-
hisce into two valves. They also fall away from the sutures,
or from a persistent marginal border of them, or in some cases
the valves thus fall away entire. The persistent frame which
remains has been called a Repium, an architectural word, here
taken in the sense of door-case.

559. A Capsule is the pod, or dehiscent fruit, of any compound
pistil. When regularly and com-
pletely dehiscent, as already stated
(544), the pod splits lengthwise into
pieces or valves. ‘The modes of regular
dehiscence are illus-
trated in Fig. 612-
617. Two modifica-
tions of the capsule
have received distinc-
tive names which are
in common use, viz.
the Pyxis and the
Silique.

560. A Pyxis or Pyxidium is a dry fruit which opens by a
circular line, cutting off the upper part as a lid; @. ef the dehiscence
is circumscisstle. (553, Fig. 621.) In the Purslane,
Pimpernel, Henbane, and Plantain, the pyxis is
a capsule; in Amaranths (Fig. 637) it is a
utricle ; in Jeffersonia (Fig. 622) it is a modi-
fication of the follicle, being of one carpel which
dehisces transversely, and not all round, so that
the lid remains attached. on

561. A Silique is a narrow two-valved capsule, with two pari-
etal placentze, from which the valves separate in dehiscence ; as
in plants of the Cruciferous or Mustard family (Fig. 623),
to the fruit of which this term is restricted. Usually, a false
partition is stretched across between the two placente, render-

FIG. 621. Pyxis of Purslane, Portulaca oleracea, the top separating entirely and
falling away.

FIG. 622. Pyxis-like follicular fruit of Jeffersonia diphylla; the lid remaining
attached dorsally. .

FIG. 623. Silique of Cardamine in dehiscence. 624. Silicle of Capsella or Shepherd’s-
Purse, lateral view, and an oblique view of the same with one valve removed.

294 THE FRUIT.

ing the pod two-celled in an anomalous manner. A SILIcLe
(Silieula, diminutive of siliqua) is merely a short silique, the
length of which does not more than twice or thrice surpass the
breadth ; such as that of Shepherd’s-Purse (Fig. 624). and of
Lunaria, Candytuft, &ce.

062. Indehiscent Dry Fruits are almost always one-seeded or
very few-seeded. If numerous, the seeds thus placed would not
be dispersed. The ordinary kinds are strictly one-
seeded, and in common language are often con-
founded with seeds. The ways in which such fruits
are dispersed are various. In the following case,
the adaptation of the pericarp to dispersion by wind
distinguishes the species of fruit.

965. The Samara, sometimes called in English a
na is an indehiscent one-seeded fruit provided
with a wing. In the White Ash
the wing is terminal (Fig. 625) ;
in other species the whole fruit
is wing-margined ; in Birch and
Elm (Fig. 626) the wing sur-
rounds the body of the pericarp ;
and the Maple fruit is a double
samara or pair of such fruits, con-

626 spicuously winged from the apex. 2

564. Akene (Lat. Achenium) is a general name for all the
une-seeded, dry and hard, indehiscent and seed-like small fruits,
such as are popularly taken for naked
seeds. But that they are true pistils,
or ovaries ripened, is evident from the
style or stigma they bear, or from the
sear left by its fall; and a section
brings to view the seed within, provi-
ded with its own proper integuments.
The name has been restricted to the seed-like fruits of simple
pistils, such as those of the Buttercup (Fig. 628,629), Anemone,
Clematis, and Geum. The style in some species of the latter
remains on the fruit as a long and feathery tail, in others as a
short and hooked one, both being agents of dissemination. The
grains of the strawberry (Fig. 653) are also akenes. ‘The name
is extended to all one-celled seed-like fruits resulting from a

FIG. 625. Samara or key of White Ash, Fraxinus Americana. 626. That of White
Elm. Ulmus Americana. 627. Double samara of Red Maple, Acer rubrum.

FIG. 628. Achenium of a common Buttercup. 629. Vertical section, showing the
seed within.

ITS KINDS. 295

compound ovary, and even when invested with an adnate calyx-
tube. Of the latter is the fruit of Composite. (Fig. 630-635.)
Here the tube of the calyx is incorpo-

rated with the surface of the ovary ;

and its limb or border, obsolete in some \y4/ |) /\/4jy
cases (Fig. 630), in others appears Wh ily
as a crown or cup (Fig. 651), or set of Man i

\ Nh
teeth or of scales (Fig. 652, 633), or as \ |
a tuft of bristles or hairs (Fig. 634, \

634

632

635), &e., called the Pappus. In the Lettuce and Dandelion
(Fig. 635), the achenium is rastrate, or beaked, 7. e. its summit
is extended into a slender beak. An akene with adnate calyx
has been termed a Cypse.a.

565. The Utricle 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, discharging the seed. In the true Amaranths,
the utricle opens by a circular line, and the upper
part falls as a lid, converting the fruit into a small
pyxis (560), —a transition form. (Fig. 637.)

566. A Caryopsis or Grain differs from the utricle
or akene in having the seed completely filling the
cell, and its thin coat firmly consolidated throughout
with the very thin pericarp; as in wheat, Indian
corn, and all other cereal grains. Of all fruits this :
is the kind most likely to be mistaken for a seed. me

567. A Nut is a hard, one-celled and one-seeded, indehiscent
fruit, like an achenium, but larger, and usually produced from
an ovary of two or more cells with one or more ovules in each,
all but a single ovule and cell haying disappeared during its
growth (536) ; as in the Hazel, Beech, Oak (Fig. 638), Chest-

FIG. 630. Achenium of Mayweed (no pappus). 631. That of Cichory (its pappus a
shallow cup). 632. Of Sunflower (pappus of two deciduous seales). 633. Of Sneezeweed
(Helenium), with its pappus of five scales. 634. Of Sow-Thistle, with its pappus of
delicate downy hairs. 635. Of the Dandelion, tapering below the pappus into a
long beak.

FIG. 636. Utricle of Chenopodium album, or common Goosefoot. 637. Utricle of an
Amaranth, by transverse dehiscence becoming a pyxis.

296 THE FRUIT.

nut, and the like. The nut is often enclosed or surrounded by
a kind of involucre, termed a Cupule; such as the cup at
the base of the acorn, the bur of the chestnut, and the leaf-
like covering of the hazel-nut. The name Glans
(sometimes Gland in English) is technically applied
to such nuts, this being their classical Latin name.

563. The fruit of the Walnuts and Hickory is
apparently a kind of drupaceous nut, or something
intermediate between a stone fruit and a nut. But
certain monstrosities give reason for supposing that
the seeming exocarp (041), which in Hickory
hardens and at maturity dehisces*in four valves,
is of the nature of an adnate involucre. The cocoanut is a sort
of fibro-drupaceous nut.

569. Nutlet, or in Latin form Nucute (Nucula), is sometimes
superfluously employed in a literal sense, as a diminutive nut.?
Of late it has acquired a good and fairly legitimate use as the
name of the seed-like, or rather akene-like, closed parts or lobes,
of crustaceous or other hard texture, into which certain bilocular
or plurilocular pericarps separate at maturity, 7. e. for the seg-
ments of a schizocarp, 571, which resemble akenes.? These are
sometimes carpels, sometimes half-carpels, as in Verbena, also
in Borraginaceee and Labiatee (in which the segments are greatly
separated in the ovary), and sometimes, as in Nolana, they are
portions of compounded carpels which have been exceedingly
multiplied by chorisis.

570. There are complete transitions between dry nutlets, with
a thin and herbaceous epicarp, and the pyrene (574) or stony
inner portion of such carpels when drupaceous or composing a
drupe of two or more stones. It is therefore a hardly incongru-
ous and very convenient use which extends the term nutlet to
include these small seed-like stones also, as, for example, to
those of Holly, Bearberry, Hawthorn, and the like.

71. The pair of achenium-like or often samara-like carpels,

1 Nut and akene, between which there is no fixed distinction, will cover
this ground. The fruit of Cyperaceex, for instance, is truly an achenium,
if this name is ever to be used (and it now commonly is) for any other than
a monocarpellary fruit. It is often termed a nut, sometimes a nutlet, and
by a late writer, Beeckler, a caryopsis.

2 Cocci (sing. Coccus, from a Greek word for kernel) is another name for
fruit-carpels, or separating lobes of a dry pericarp, as well for dehiscent ones
(of Euphorbia) as for indehiscent. Hence such lobed or partible fruits
are said to be dicoccous, tricoccous, &e., according to the number of lobes or
carpels.

FIG. 638. Acorn (nut) of White Oak, with its cup. or enpule.

2-

ITS KINDS. 297

united by their inner face but separating entire at maturity,
which constitute the fruit of Umbellifersze, takes the name of
Cremocarp (Lat. Cremocarpium); and the halves are called
Mericarps. These names it may sometimes be convenient to
use; yet it is not advisable to have special names for the fruits
of particular families; and mericarp is here synonymous with
earpel. For dry fruits in general (or such as become dry)
which are composed of two or more carpels, and which at matu-
rity split up or otherwise separate into two or more closed one-
seeded portions, an appropriate recent name is that of Scuizocare.
The component carpels of such a fruit were long ago named Car-
cerules (carceruli, little prisons) by Mirbel.

572. Fleshy Fruits, which from their texture are naturally
indehiscent, may be either fleshy throughout, or with a firm rind
or shell, or fleshy externally and hard or stony internally. Of
the latter, the type is

075. The Drupe or Stone Fruit proper (Fig. 639), that of the
cherry, plum, and peach. ‘True drupes are of a single carpel,
one-celled and one-seeded
(or at most two-seeded), in
the ripening of which the
outer portion of the pericarp
becomes fleshy or pulpy, and
the inner stony or crustace-
ous, 7. é. divides into sarco-
carp and putamen. (541.)
But the name is extended to oe at
pericarps of similar texture resulting from a compound pistil,
either of a single cell, as in Celtis, and (by abortion) in the olive,
or of two or several cells, as in Cornus, Rhamnus, &c. The several
pericarps of the aggregate blackberry and raspberry are diminu-
tive drupes or DruPe.ets.

574. Small drupes are often confounded with berries, and the
stone or stones taken for seeds. Especially is it so in drupes
or drupaceous fruits of more than one cell, ripening into separate
or separable hard endocarps or stones, each filled by a seed.1
Bearberries (Arctostaphylos) and Huckleberries (Gaylussacia)
are good illustrations of this. The seed-like endocarps of this

1 The term Ac/inus, the original name of such a berry as a grape, has been
used in descriptive botany for a small drupe or drupelet, and the ripened
carpels of Rubus have been termed acini or acines, but without discriminating
them from berries.

FIG. 639. Vertical section of a peach. 640. An almond; in which the exocarp, the

portion of the pericarp that represents the pulp of the peach, remains juiceless, and at
length separates by dehiscence from the endocarp, or shell.

298 THE FRUIT.

sort are PrrEN&; and the fruits are dipyrenous, tripyrenous,
tetrapyrenous, &c., according as they contain two, three, or four
pyrene. When the sarcocarp is thin and dries up at maturity,
these pyrenz pass by gradations into nuculz (569) or nutlets:
hence pyrene are not uncommonly in English descriptions called
nutlets or nucules.

575. The Pome (Fig. 641, 642) is the name of the apple, pear,
and quince. These are fleshy fruits, composed of two to
several carpels (rarely by abortion only one), of
parchment-like or (in Hawthorns) bony texture,
enclosed in flesh which morphologically belongs
to adnate calyx and receptacle; as may be ap-
prehended by comparing a rose-hip (Fig. 407, in
flower) with an apple or a pear. Of the quince, the
whole flesh is calyx or hypanthium (395) ; in the
apple and pear, the inner or core-portion of the
flesh is of the nature of disk, investing the carpels.
In the fruit of Hawthorns, the carpels become bony
pyrene (074), and so the fruit is drupaceous, is
indeed nothing more than a syncarpous drupe.
In Eriobotrya, or Cumquat, the carpels becoming
very thin and membranaceous, the pomaceous
fruit is in fact a kind of berry.

576. The Pepo, or Gourd-fruit (Fig. 643), of which the gourd
and squash are the type, and the melon and cucumber equally
familiar illustrations, is the char-
acteristic fruit of Cucurbitacez,
fleshy internally and with a hard
or firm rind, all or part of which
is referable to the adnate calyx
completely incorporate with the
ovary. This is either one-celled
with three broad and revolute.
parietal placente, or these pla-
cent, borne on thin dissepiments,
meet in the axis, enlarge, and
spread, unite with their fellows

‘ on each side, and are reflected to
the walls of the pericarp, next which they bear their ovules. As
the fruit enkarges, the seed- -bearing placentse usually cohere
with the walls, and the partitions are obliterated, giving the

FIG. 641. Pome or apple in transverse section. 642. Quince in vertical section: the
inner flesh answering to disk in the apple and pear is here wanting.

FIG, 643. Section of the ovary of the Gourd. 644. Diagram of one of iis constituent
earpels.

ITS. KINDS. 299

appearance of a peculiar abnormal placentation, which the study
of the ovary readily explains. In the watermelon the edible
pulp all belongs to the greatly developed placente. Fruits of
this family in which the rind also is soft at maturity are true
berries.

577. The Hesperidium (orange, lemon, and lime) is the fleshy
fruit of a free many-celled ovary with a leathery rind, and is a
mere variety of the berry. The name is
applied only to fruits of the Orange tribe.

578. The Berry (Lat. Bacca) comprises
all simple fruits in which the pericarp is
fleshy throughout. The grape, gooseberry,
currant, cranberry (lig. 645), banana, and
tomato are familiar examples. The first
and last consist of an ovary free from the
calyx ; in the others, calyx and ovary are
combined by adnation.

579. Aggregate Fruits are those in which
a cluster of carpels, all belonging to one
flower, are crowded on the receptacle into
one mass, as in the raspberry and _ black-
berry taken as a whole. (Fig. 646.) They C7
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 akenes, 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 Syxcarprom, or in English
form Syncarp. But the term has
been applied to multiple fruits as
well.! In Hydrastis, the numerous |
earpels imbricated on the upper
part of the torus are baccate, that
is, become berries ; in araspberry, 6i7
the seemingly baccate grains are drupaceous (being drupelets, 573),

1 The syncarp which is a gynecium might be designated a simple syn-
carpium ; that which is an inflorescence, a complex syncarpium, which may be
biflorous, pauciflorous, or multiforous.

FIG. 645. The larger Cranberry, Vaccinium (Oxycoccus) macrocarpon; the one
transversely divided.

FIG. 646. Vertical section of half of a blackberry (of Rubus villosus), enlarged; and,
647, of one of its drupelets more magnified.

300 THE FRUIT.

and, slightly cohering together (though without organic union),
they fall as one body from the conical dry torus at maturity. It
is the same in blackberries or bramble-berries (Fig. 646, 647),
except that the drupelets persist on the torus, which partakes of
the juiciness. In the aggregate fruit of Magnolia (Fig. 648-650),
such carpeis, imbricated over one another, cohere more or less
at all contiguous parts, and
become drupaceous ; neyer-
theless, at maturity each
opens dorsally, allowing the
seeds to fall out: in age it
dries and hardens, and also
separates from its conneec-
tions, and so_be-
comes a follicle, but
with the remark-
able peculiarity of
dorsal instead of
ventral dehiscence.
(Fig. 650.) In Li-
riodendron, a tree
of the same family,
such carpels are
dry and indehiscent throughout ; and they largely consist of long
and flat styles, imbricated in. a cone, but separating from each
other and from the slender torus at maturity, when each becomes
a samara.

580. Accessory or Anthocarpous Fruits are those of which sq@me
conspicuous portion of the fructification neither belorigs to the
pistil nor is organically) united with it, except by a common
insertion. The part thus imitating a fruit, while it is really no
part of the pericarp, is sometimes called a Pseudocarp, or an
Anthocarp or Anthocarpium. This condition may occur either
in simple, in aggregate, or in multiple fruits.

1 The aggregate fruit like that of Rubus (named by some Conocarpium,
by others an -Lterio, Erythrostomum, &c.) was termed by Dumortier a Drupe-
tum. <A similar aggregation of baccate carpels he termed a Baccetum ; of
follicles, a Follicetum, &¢. All such names may look well ina system; but
they are both superfluous and unmanageable in phytography.

FIG. 648. Aggregate fruit of Umbrella-tree, Magnolia Umbrella, reduced in size; a
seed from a lower dehiscent carpel hangs on « thread. consisting of a tuft of extensile
spiral ducts unravelled. 649. Same in longitudinal section. 650. One of the carpels
detached, at full maturity, dried up, dorsally dehiscent, exposing the pair of seeds of
the natural size. :

ITS KINDS. 301

5381. Gaultheria procumbens, the aromatic Wintergreen (Fig.
651, 652), affords a good example of the first. Its seeming
berry (the checkerberry), with summit crowned by the tips of
the calyx-lobes, well imitates the true
berry of a Vaccinium, such as that of
Fig. 645. But it comes from a flower
with thin calyx, underneath and free
from the ovary. Its fruit is really
a capsule: in the process of fructi-
fication, the calyx enlarges, becomes
succulent, completely encloses the capsule or true fruit, yet
without adhering to it, and in ripening counterfeits a red
berry. So in Shepherdia, or Buffalo Berry, the seeming sarco-
carp of a drupe is really a free calyx, accrescent and succulent,
enclosing an akene. So, also, the apparent achenium or nut of
Mirabilis, or Four-o’clock, and of its allies,
is the thickened and indurated base of the
tube of a free 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 (889, Fig. 406, 655), the akenes or
true fruits dispersed 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 combined, as in a pinc-apple.

583. Multiple or Collective Fruits + are those which result from .
the aggregation of several flowers into one mass. The simplest
of these are those of the Partridge-Berry (Mitchella, Fig. 467),

bol

65383

1 Collective is the preferable name. The term mu/tiple was applied by
DeCandolle to what are here (following Lindley) called aggregate fruits ;
‘and the aggregate fruits of DeCandolle are here called multiple or collective.
Moreover, the distinction between accessory or anthocarpous and collective
or multiple fruits was not recognized by Lindley, who combined the two —
in his original “Introduction to Botany.” In this work four classes are
given: 1. Fruit simple, Apocarpi; 2. Fruit aggregate, AGGREGATI;

FIG. 651. Forming capsule of Gaultheria procumbeps, with enlarging calyx partly
covering it. 652. Same, more advanced, ani in longitudinal section.
FIG. 653. Vertical section of half a strawberry. Compare with Fig. 406.

302 THE FRUIT.

and of certain species of Honeysuckle, formed of the ovaries
of two blossoms united into one fleshy fruit. The more usual
sorts are such as the pine-apple, mulberry, and the fig. These
are, in fact, dense forms of inflorescence, with the fruits or floral
envelopes matted together or coherent with each other; and all
or some of the parts succulent. The grains of the mulberry
(Fig. 654-656) are not the ovaries of a singie flower, like those

659 655 656

of the blackberry, which it superficially resembles: they belong
to as many separate flowers ; and the pulp pertains to the calyx,
not to the pericarp, which is an akene. So that this, like most
multiple fruits, is anthocarpous as Well as multiple. Similarly,
the mostly indefinite fructiferous masses of Strawbgrry Blite may
resemble strawberries ; but the pulpy part is the Calyx of many
flowers, not the succulent receptacle of one. In the pine-apple,
the flowers are spicate or capitate on a simple axis, which grows
on beyond them into leafy stem; this when rooted as a cutting

3. Fruit compound (ovaria compound), Syncarpr1; 4. Collective fruits,
Anthocarpi. .

Later, in his “ Elements of Botany,” Lindley reduced the classes to
two: 1. Simple Fruits, those proceeding from a single flower; 2. Multiple
Jruits, those formed out of several flowers.

FIG. 654. A mulberry. young. 655. One of the fleshy grains at flowering time, show-
ing it to be a pistillate blossom with fleshy calyx. 656. Thesame later, with the succu-
lent sepals in transverse section.

FIG. 657. A young fig. 658. Longitudinal section of the same later, but in flowering
time. 659. A small slice, magnified, showing some of the flowers.

ITS KINDS. 303

bears another pine-apple, and so on: the constituent flowers have
through immemorial propagation in this way become sterile and
seedless, and all its parts, along with the bracts and the axis of
the stem, blend in ripening into one fleshy and juicy mass. Few
fruits of this class have ever been technically named, at least
with names which have come into use. But the two following
deserve special appellations, although only the latter is familar
either in ordinary language or in descriptive botany. ;
584. The Syconium or Hypanthodium, the Fig fruit. (ig. 657—
659.) This results from a multitude of flowers concealed in a
hollow flower-stalk, if it may be so called, which becomes pulpy
and edible when ripe; and thus the fruct 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
Mito very small akenes, which are commonly taken for seeds.
The fig is to the mulberry what a rose-hip is to a strawberry.
(389, Fig. 406, 407.) It is further explained by a comparison
with a near relative of the Fig-tree, Dorstenia, in which similar
flowers cover the upper surface of a flat peltate disk. This disk
or plate sometimes becomes saucer-shaped by. an elevation or
incurvation of the margin. <A greater degree of this would
render it cup-shaped, or even pitcher-shaped ; from which it is
a short step to the contraction of the mouth down to the small
orifice which is found in the fir. ;
_ 585. The Strobile or Cone (Fig. 660) is a scaly multiple fruit,
resulting from the ripening of certain sorts of catkin. The name
is applied to the fruit of the Hop, where
the large and thin scales are bracts ;
but it more especially beiongs to the
Pine or Fir cone, the peculiar fruit of
Coniferze (507), in which naked seeds
are borne on the upper face of each
fructiferous scale (Fig. 661), or gome-
times in their axils.
Such a cone when
spherical, and of
thickened scales
with narrow base, as
that of Cypresses,
has been termed a
GALBULUs, an unnecessary name. The galbulus of Juniper is a

FIG. 660. Strobile or Cone of a Pitch Pine, Pinus rigida. 661. Inside view of one
of the scales, showing one of the winged seeds, and the place from which the other, 662,
has been detached.

304 THE FRUIT.

remarkable transformation into a seeming berry; the few scales
cohering with each other as they grow and becoming fleshy at
maturity, completely enclosing a few bony-coated seeds.

586. A Synopsis of the kinds of Fruit, as characterized in
this chapter, is appended. The analysis extends only to simple
fruits. For there are no commonly used special names of
kinds of Aggregate (579), Accessory (580), or Multiple (583)
fruits, except that of Strobile.

SIMPLE FRUITS are

Dry and dehiscent, monocarpellary,

Opening by one (chiefly the ventral) suture, . . . . . . . FOLLicte.
Opening by both sutures;;.-: 2 2. 2 2°90) & 2°) See
Or transversély jointed; 2) 22. 1 3) 2
Dry and dehiscent, bi-pluri-carpellary,. . . . . + . . . « CAPSULE.
When its dehiscence is circumscissile, . . .- oJ. «= gJPyoge
When dehiscent by two valves from two eel een. . SILIQUE.
A short and broad silique,. . . : 3 32 SEECrEE
Dry and bi-pluri-carpellary, splitting = Gneseedea carpal . SCHIZOCARP.
The dimerous schizocarp of Umbellifere, . . . . . . .CREMOCARP.
Each of its halves or carpels,. . - . . . Hemicarp or MERICARP.
The akenelike or nut-like Pe into which Schizocarps generally
Mivides sa.) 0 . . Nucures or NUTLETS.
Dry and indehiscent, queeetet one swordceded!
Winged} 2.04 eer MSS OMe hfe os See EL
Wingless, and with the
Thin pericarp consolidated with the seed, . . . . . . Caryopsis
Thin pericarp loose and not filled by the seed, . . . . . Urricie.

Thick or hard pericarp free from the seed,
Small, from a one-celled one-two-ovuled ovary, AKENE or ACHENIUM.

Larger, mostly from a two-several-celled and ovuled ovary, . Nut.
Nut borne in a cupule orinvolucre, ... ... . . . GLANS.

Fleshy and indehiscent,

Heterogeneous in texture, having
A stone (putamen) or nutlets within an exterior sarcocarp, . DRuPE.
Papery or cartilaginous carpels in an inferior sarcocarp, . . Pome.

A harder or firm rind or exterior, and soft interior,

From an inferior ovary (confined to Gourd Family), . . . Prpo.
From a superior ovary (confined to Orange Family), Hrsperrpicum.
Homogeneous, fleshy throughout, .... . . . + + « «+ BERRY.

THE SEED. 305

CHAPTER VII.
THE SEED.

507. Tue Seep is the fertilized ovule (515), with embryo
formeG within it. It consists, like the ovule, of a nucleus or
kernel, enclosed by integuments. The seed-coats are those of
the ovule, viz. two, or sometimes only one, in certain plants
none. Occasionally an accessory coat appears after fertiliza-
tion; and certain appendages may be produced, as outgrowths
from some part of its surface or from its base. The nucleus or
kernel is composed either of the embryo alone, or of a nutritive
deposit in addition. (19-41.) All the parts of a seed are in-
dicated in Fig. 665.

588. The SrEep-staLK or PoposPERM, when there is one, is
the funiculus of the ovule (516), and retains this name. So
also do the Cuaraza, Ruaprue, and Hitum; the latter being the
scar left by the separation of the seed from a
its funiculus or directly from the placenta.
The foramen of the ovule, now closed, is
the MicropyLe of the seed.

589. The terms which denote the char-
acter of the ovule, such as orthotropous,
campylotropous, amphitropous, and anatropous, apply equally to
the resulting seed.

590. Seed-Coats. The integuments of the seed answer to the
primine and secundine of the ovule. The main seed-coat is the
exterior integument of the ovule when there is more than one.
Being the most firm coat, and not rarely crustaceous in texture,
it takes the name of Trsra, which is equivalent to seed-shell.
It has also been named Spermoperm (seed-skin). and sometimes
Episperm. The latter name (meaning upon the seed) is best
applied to the pellicle or outer layer, sometimes a thick one,
which the testa of certain seeds forms. The testa is extremely
various in form and texture, is either close and conformed to

FIG. 663. Vertical magnified section of the (anatropous) seed of the American Lin-
den; with the parts indicated, viz. the hilum (a); testa (5); tegmen (c); albumen (4);
embryo (¢). 664. Vertical section of the orthotropous seed of Helianthemum Cana-
dense, with its funiculus, a.

306 THE SEED.

the nucleus, or loose and cellular (as in Pyrola-seeds), or vari-
ously appendaged.

591. The inner coat, ~1lled TreGmMeN and sometimes Enpo-
PLEURA, When present is always conformed to the nucleus, and
is thin or soft and delicate. Sometimes it is inconspicuous
through cohesion with the nucleus or with the inner surface of
the testa. In ovules of one coat it is necessarily wanting.

592. Appendages or outgrowths of the testa generally have
reference to dissemination. Iwo characteristic kinds of such
appendages are the wing and the coma, -
both pertaining only to the seeds of dehis-
cent fruits and calculated, by rendering
seeds buoyant, to facilitate dispersion by
the wind. ‘The wing of a Pine-seed (Fig.
661, 662) is a part of the carpellary scale
upon which the two ovules grew. In
Trumpet Creeper (Fig. 665), an entire
wing surrounds the body of the seed.
In the related Catalpa (Fig. 666), it is
mainly extended from the two ends, and
almost dissolved into a coma, the name
given to the tuft of soft hairs like that
which forms the down at one end of the
seed of Milkweed (Fig. 667), and of
Epilobium, and at both ends in several
Apocynaceze. In the Cotton-plant, very
long and soft hairs, admirably adapted for
spinning, thickly cover the whole seed-

65 = coat. The wing and coma of seeds are
functionally identical with the wing and the pappus of the pericarp
in the samara and the akenes of Composite (563, 564), but
morphologically quite unlike them.

593. There are other (mainly microscopic) structures on some
seed-coats which come usefully into play in arresting farther
dispersion at a propitious time or place. In many but not all
Polemoniacez (notably in Collomia), in certain Acanthacee,
such as Ruellia tuberosa (and equally in certain Composite of
the Senecio tribe and in Salvias, &c., among Labiatee, where
this structure is transferred to akenes and nutlets), the testa is
coated with short hairs. which when wetted burst or otherwise
open and discharge along with mucilage one or more very atten-

FIG. 665. Winged seed of Trumpet Creeper, Tecoma radicans. 666 That of Catalpa,
becoming comose: the body divided lengtliwise through the embryo.
FIG. 667. Comose sced of Milkweed, Asclepias Cornuti.

ITS COATS AND APPENDAGES. 307

uated long threads (spiricles) which were coiled within. These,
protruding in all directions and in immense numbers, form a
limbus of considerable size around the seed, and evidently must
serve a useful end in fixing these sma i
and light seeds to the soil in time of
rain, or to moist ground, favorable to
germination. In cress and flax-seed, i
the abundant mucilage developed when a——)-
wetted comes from the gelatination of
epidermal cell-walls, and
subserves a similar use.
.094. While the testa in
many seeds is hard and
crustaceous or bony, imitat- 668 64)
ing the pericarp of a nut, in others (such as Peonia) it becomes
berry-like (baccate), and in Magnolia, drupaceous.’ (Fig. 668-
671.) These may also be regarded as adaptations for dissemi-

67

nation, here by the agency of birds, attracted by bright coloring
and edible pulp.

595. The rhaphe of an anatropous seed (shown in Fig. 681,
685) is sometimes so salient as to form a conspicuous appen-
dage, as in Sarracenia, Fig. 672. Again it may be wholly

1 See article On the Structure of the Ovule and Seed-coats of Magnolia,
in Jour. Linn. Soc. ii. 106, from which the accompanying figures and Tig.
589-597 are reproduced.

FIG. 668. Forming seed (one eighth of an inch long) of Magnolia Umbrella; the
rhaphe toward the eye. 669. Magnified view of the same divided lengthwise through
the rhaphe; the outer coat, a, beginning to form a hard inner layer, a’. Within and
distinct from this is the inner coat (b), immediately enclosing the nucleus, c. The oppo-
site side of the testa is thicker on account of the rhaphe, in which d indicates the cord
of spiral ducts.

FiG. 670. A nearly full-grown seed, of the natural size. 671. Longitudinal section,
emiargen, suowing the crustaceous or stony inner stratum of the testa well developed;
the parts lettered as in Fig. 669. 672. A transverse section in the same position.

308 THE SEED.

inconspicuous, as in the ripe seed of Magnolia, where it is at
length completely merged and imbedded in the fleshy drupaceous
testa, as shown in Fig. 670-672.

596. Crest-like or other appendages are not uncommon either
on the rhaphe or at the hilum. These are outgrowths produced
during the development of the ovule into the seed. In Sangui-
naria, such a crest develops from the whole length of the rhaphe

672 675 674 65

(Fig. 673) : in Dicentra, Corydalis (Fig. 674), &c., from some
part of it, mostly from its base next the hilum, or from the
hilum itself, or even from just below it. Such an appendage,
especially when attached to the base of the seed, is named a
StROpHIOLE. A similar and commonly a wart-shaped appendage
in Euphorbia, Ricinus (Fig. 675), &e., is produced by an out-
growth of the external orifice of the ovule, the micropyle of the
seed. This properly takes the name of Caruncie. But the
two terms are not always discriminated. By further develop-
ment, either of these may give rise, in certain seeds, to an acces-
sory covering called

097. The Aril or Arillus. This term, rather vaguely employed
by Linnzeus, was first well defined by Geertner. The true arillus
is an accessory seed-covering, more or less incom-
plete, formed between the time of fertilization and
the ripening of the seed, by a growth from the apex
of the funiculus (when there is any) at or just be-
low the hilum, in a manner similar to that in which
the coat or coats of the ovule are formed. That
of Nymphiea (Fig. 676) is a typical example ; only
the arillus is developed from the funiculus at a point distinctly
below its apex: here a ring forms, which grows into a cup, and
this is soon extended into a sac, loosely enclosing the seed, and
open at the top. This is membranaceous : commonly it is fleshy.
When there is absolutely no funiculus. the aril may originate
from the placenta, as it does in Podophyllum, in which most of

FIG. 672. Anatropous seed of Sarracenia purpurea, with very salient rhaphe. 673.
Same of Sanguinaria or Bloodroot, with rhaphe crested for its whole length 674. Seed
of Corydalis aurea, with crest or strophiole, attached at or near the hilum. 675. Seed
(suspended) of Ricinus, with its caruncle.

FIG. 676. Seed of White Water-Lily, Nymphaa odorata, in its loose and thin arillus.

~)

ARILLUS, ALBUMEN. 300

S,

the pulp of the berry consists of these fleshy arils, much com-
pacted. (Fig. 677, 678.)

598. The laciniate aril of the nutmeg (mace) and, it is said,
the bright red and pulpy aril of Euonymus and Celastrus begin
in the manner of a ca-
runcle, and are formed
(mainly if not wholly)
of an outgrowth at or
around the micropyle. So
that, if an orthotropous
seed ever deyeloped an
aril of this sort, it would 677 673
be seen to begin at the apex of the seed and cover it from above
downward. Planchon, who distinguished this from the true aril,
gave to it the name of ArILLODE (Arillodium) or False Arillus.

599. The Nucleus, or kernel of the seed, consists of the Albu-
men, when this substance is present, and the Embryo.

600. The Albumen, as described in the second chapter (25, &e.),
is the name generally employed by systematic botanists for a
store of nutritive matter in the seed outside of the embryo,
whatever its chemical composition. It is not here the name
of a chemical substance (albumen or albumin), but of a cellular
structure, the cells of which are loaded commonly with starch-
grains (as in the Cerealia), more or less mingled with other
matters, or else filled with an encrusting deposit of some equiva-
lent substance, as in the cocoanut, coffee-grain, &e. The cells
in which this deposit is made 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 stage. In what are called albuminous seeds it persists and
forms either almost the whole kernel, the embryo remaining
minute (as in Fig. 23,54, 680), or forms a large portion of it (Fig.
13, 17, 19, 21, 48, 663, 664), or, by the growth of the embryo
displacing it, it may in the ripe seed be reduced to a thin stratum
or mere lining to the contiguous seed-coat; or it may disappear
altogether, as in the seeds of Maple, Almond, Squash, Pea, and
the like, which are therefore said to be exalbuminous. The
difference between albuminous and exalbuminous seeds is that
the maternal nutritive deposit is transferred to the embryo in

FIG. 677. Section of pericarp and placenta of Podophyllum peltatum; the pulp of
the latter mainly of the nature of arillus. investing the seeds. 678. The arillus of ong
seed detached and enlarged, divided lengthwise, showing the seed within.

310 THE SEED.

the former during germination, in the latter during the growth
of the seed.

602. The albumen was named Perisperm by Jussieu, and
Endosperm by Richard (25, note); but neither name has in
systematic botany displaced the earlier one of Grew and Geert-
ner. But both names have recently been brought into use to
distinguish between two kinds of albumen, that formed within
the embryo-sac, which is specifically termed ENpospErmM, and that
formed without, which takes the name of PEerisperM. ‘This use
comports with the etymology of the two words, the former refer-
ring to a comparatively internal and the latter to an external
portion of the seed or kernel.

603. In most seeds the albumen is endosperm: in Canna it
is all perisperm. In Nymphvea and its allies (except Nelum-
bium, which has none) mostof it is perisperm ;
but a thin and condensed layer of endosperm
surrounds the embryo, where with the per-
sistent embryo-sac (or the apex of it) it
forms the fleshy sac in which the embryo is
enclosed. It is the same in the Pepper Family
(Fig. 679), except that there is a larger quan-

679 tity of endosperm or inner albumen.,

604. When the nucleus of a ripe seed is hollow, as in the
cocoanut and nux vomica, the formation of endosperm, which
usually begins next the wall of the embryo-sac, has not proceeded
so as to fill the cavity. The embryo-sac in the cocoanut attains
enormous size, and the cavity is filled by the milky fluid.

605. The texture or consistence of the albumen differs greatly.
It is fartnaceous or mealy when, consisting mainly of stareh-
grains, it may readily be broken down into a powder,
as in wheat, buckwheat, &c. ; o¢/y, when saturated
with a fixed oil, as in poppy-seed; fleshy, when
more compact, but readily cut with a knife, as in
the seed of Barberry ; muctlaginous, when soft and
somewhat pulpy, as in Morning Glory and Mallow,
but when dry it becomes fleshy or harder ; corneous,

6S) when of the texture of horn, as in coffee and the
seed of Caulophyllum ; and even ony, as in the vegetable ivory,
the seed of Phytelephas. It is mostly uniform ; but in the nutmeg,

FIG. 679. Longitudinal magnified section of a seed of Black Pepper; showing the
large episperm, the small endosperm in the persistent embryo-sac, and in this the
minute embryo.

FIG. 680. Longitudinal section of a seed of the so-called Papaw, Asimina triloba,
with ruminated albumen and minute embryo.

THE EMBRYO. Sit

in the seeds of Asimina (Fig. 680) and all the Custard-Apple
family, it is marked by transverse lines or divisions (caused by
inflexions or growths of the inner seed-coat), giving a section
of it either a marbled appearance, or as if it had been slit by
incisions: it is then said to be ruménated.

632 Got

606. The Embryo,’ being an initial plantlet or individual of a
new generation, is of course the most important part of the seed.
To its production, protection, and support, all the other parts of
the fruit and flower are subservient.

607. In an embryo of full development, namely, one in which
all the parts are manifest antecedent to germination, these parts
are the Caulicle, otherwise called Radicle, the Cotyledons, and
the Plumule. (20, 30.) The first is the initial
axis or stem, a primary internode; the second
consists of the leaves of the primary node; the
third is a beginning of a farther growth which
is to develop more stem and leaves. Such an
embryo is usually unaccompanied by albumen,
having in the course of its growth taken into
itself (mostly into the cotyledons) the provision
which in other seeds is mainly accumulated external to it until it
is drawn upon in germination.

1 The word Embryo or Embryon was applied to this body in plants by
Bonnet (Considérations sur les Corps organisées), in 1762, and was introduced
into systematic botany at about the same time (1763) by Adanson: it was
taken up by Gertner in 1788. Jussieu in the Genera Plantarum (1789) held
to the term Corculum (the cor seminis) which came down from Cesalpinus.

Being the germinal part of the seed, the embryo of the plant, like that of
the animal, is in general language often called the Germ.

FIG. 681. Seed of a Violet (anatropous), enlarged; with hilum or scar (a), rhaphe
(6), and chalaza (ce) indicated. 682. Vertical section of the same, showing the straight
embryo in the axis of the mealy albumen.

FIG. 683. Vertical section of the (orthotropous) seed of Buckwheat, showing the
embryo folded round in the mealy albumen.

FIG. 684. Vertical section of the (anatropous) seed of Elodea Virginica, the embryo
completely filling the coats

FIG. 685. Seed of Delphinium tricorne (anatropous), enlarged; the hilum, the
rhaphe. and the chalaza lettered as in Fig. 681. 686. Vertical section of the same with
ce. the chalaza, d, the testa, e, the tegmen,.f, the albumen, g, the minute embryo near
the hilum (a).

FIG. 687. Embryo of the Pumpkin, with its short radicle and large and flat cotyle-
dons, seen flatwise. 688. A vertical section of the same, viewed edgewise.

312 THE SEED.

608. The opposite extreme is an embryo (as in Fig. 686)
which appears as a mere speck in the albumen, but in which
close microscopical inspection may commonly reveal some differ-
entiation, such as a slight notch at one end (that farthest re-
moved from the micropyle) of a dicotyledonous embryo, indicating
the future cotyledons. Indeed, in Monotropex, Orobanchacee,
and some other parasitic dicotyledonous plants, and in Orchids
among the monocotyledonous, the embryo is a globular or oblong
particle, with no adumbration of organs whatever antecedent to
germination. There are all grades between the most rudimen-
tary and the most developed embryos.

609. Under the circumstances of its formation (532), the
radicular end of the embryo is always near to and points towards
the micropyle of the seed, viz. to what was the orifice of the
ovule ; and if the embryo be straight, or merely partakes of the
curvature of the seed, the cotyledons point to the opposite
extremity, that is, to the chalaza.

610. The position of the radicle as respects the hilum varies
with the different kinds of seed. In the orthotropous form, as in
Helianthemum (Fig. 664) and Pepper (Fig. 679), the radicle
necessarily points directly away from the hilum.’ In the anatro-
pous form, as in Fig. 663, 682, and 684-686, the extremity
of the radicle is brought to the immediate vicinity of the hilum;
and so it is, although in a different way,
in the campylotropous seed (Fig. 689,
690); while in the amphitropous the
radicle points away from the hilum later-
ally. As the nature of the ovule and seed
may usually be ascertained by external inspection, so the situation

1 Two technical terms, early introduced by Richard to indicate the direc-
tion of the radicle (caulicle), or rather its relation to the hilum, are

Antitropous, when the embryo directs its radicle away from the hilum, as
it must in all orthotropous seeds ;

Orthotropous, also homotropous, when directed to the hilum (more strictly to
the micropyle close to the hilum), as in anatropous seeds. These two terms
are still employed by many botanists, although superfluous when the ovule
or seed is stated to be anatropous or orthotropous, &c. And the term
orthotropous, so used, is liable to be confused with orthotropous as applied
to the ovule.

Richard, moreover, termed the embryo amphitropous when curved or coiled,
as in Chickweed (Fig. 689) and all such campylotropeus seeds; and hetero-
tropous when neither radicle nor cotyledons point to the hilum, as occurs
in the semi-anatropous or amphitropous ovule. Many botanists describe
the last by the expression “ radicle vague,” or, better, “ embryo transverse.”

FIG. 689. Campylotropous seed of common Chickweed, magnified. 690. Section ot
the same, showing the embryo coiled into a ring around the albumen.

THE EMBRYO. 313

of the embryo within, and of its parts, may often be inferred
withont dissection. But the dissection of seeds is not generally
difficult.

611. The direction of the radicle with respect to the pericarp
is also noticed by systematic writers; who employ the terms
radicle superior or ascending when this points to the apex of the
fruit; radicle inferior or descending when it points to its base ;
centripetal, when turned toward the axis of the fruit; centrifugal
(or peritropous), when turned toward the sides; and vague,
when it bears no evident or uniform relation of the kind to the
pericarp.

612. The position of the embryo as respects the albumen,
when that is present, is various. Although more commonly in
the axis, it is often excentric, or even external to the albumen,
as in all Grasses and cereal Grains (Fig. 56-61), in Polygonum,
&c. When external or nearly so, and curved circularly around
the albumen, as in Chickweed (Fig. 690) and Mirabilis (Fig.
17), it is said to be peripherie.

613. The embryo may 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 etreinately con-

GOL 692 695

volute from the apex, as in Bunias; or else doubled up and thus
biplicately convolute, as in Sugar Maple, Fig. 2. Two modi-
fications are more common, and are of such classificatory impor-
tance in Cruciferze as to need special reference. Namely, when
cotyledons are

Incumbent (as in Fig. 692, 693), being so folded that the back
of one is laid against the side of the radicle ; and

Accumbent (Fig. 694, 695), when the edges of the pair of
cotyledons are longitudinally applied to the radicle. These
differences were first employed in the classification Cruciferze by

FIG. 691. Convolute embryo of Calycanthus, the upper half cut away.

FIG. 692. Seed of a Cruciferous plant (Sisymbrium), with incumbent cotyledons,
divided. 693. Embryo of the same detached entire.

FIG. 694. Seed of a Cruciferous plant (Barbarea) with accumbent cotyledons
695. The embryo entire.

314 THE SEED.

Robert Brown, and were adopted as primary and tribal characters
by DeCandolle.

614. As to number of cotyledons, the two types of embryo
are the

Monocotyledonous, with a single cotyledon, 7. e. leaves at the
first nodes alternate (39); and

Dicotyledonous, with a pair of cotyledons, 7.e. leaves of the
first node in the most simple whorl, a pair, in other words, oppo.
site (21) ; with its modification of

Polycotyledonous (38), the leaves of the first node in whorls of
three, four, or more. This occurs with constancy in a majority
of Conifer (Fig. 48, 49), occasionally and abnormally in sundry
ordinary dicotyledonous species.

615. There are several embryos of the cotyledonous type in
which one cotyledon is smaller than the other, viz. the inner
one when the embryo is coiled or folded. And in all the species
of Abronia (a genus allied to Mirabilis, Fig. 18) this cotyledon
is wanting, so that the embryo becomes technically monocotyle-
donous. In another genus, the Dodder (Fig. 78, 79), both
cotyledons are constantly wanting ; and the plumule shows only
minute scales, the homologues of succeeding leaves reduced
almost to nothing.

616. Sometimes the two cotyledons are consolidated into one
body by the coalescence of their contiguous faces; when they
are said to be conferruminate. This occurs more or less in the
Horsechestnut and Buckeye (Fig. 41, 42), and is striking in
the seed of the Live Oak, Quercus virens.

617. The general morphology of the embryo and its develop-
ment in germination were described at the commencement of this
volume. And so the completion of this account of plant, flower,
fruit, seed, and embryo brings the history round to the starting
point. (12-19, &c.) Having mastered the morphology and
general structure of the higher grade of plants, the pupil may
go on to the morphology and structure of cells (or Vegetable
Anatomy or Histology), and to the study of Cryptogamous
Plants in all their grades.

TAXONOMY. 315

CHAPTER IX.
TAXONOMY.

Section I. Tue PrivcipLes or CLassIFICATION IN NATURAL
History.

618. Taxonomy, from two Greek words which signify arrange-
ment and law, is the study of classification. This is of utmost
importance in Natural History, on account of the vast number
of kinds to be set in order, and of relations (of agreement and
difference) to be noted. Botanical classification, when complete
and correct, will be an epitome of our knowledge of plants.
Arrangement according to kinds, and of special kinds under
the more general, is common to all subjects of study. But the
classification in Biological Natural History, that is in Botany and
Zoology, has a foundation of its own.

619. The peculiarity of plants and animals is that they exist
as individuals, propagating their like from generation to genera-
tion in a series. Of such series of individuals there are very
many kinds, and the kinds have extremely various and unequal
degrees of resemblance. There are various gradations, but not
all gradations of resemblance. Between some, the difference is
so wide that it can be said only that they belong to the same
kingdom ; between others, the resemblance is so close that it
may be questioned whether or not they came from common
parents or near ancestors.

620. The recognition of the perennial succession of similar
individuals gives the idea of Spectres. The recognition of un-
equal degrees of likeness among the species is the foundation
of GreneRA, Orpers, CLasses, and other groups of species.

621. Individuals are the units of the series which constitute
species. The idea of individuality which we recognize through-
out the animal and vegetable kingdoms is derived from ourselves,
conscious individuals, arid from our corporeal structure and
that of the higher brute animals. This structure is a whole,
from which no part can be abstracted without mutilation. Each
individual is an independent organism, of which the component
parts are reciprocally means and ends. Individuality is a main

316 TAXONOMY.

distinction between beings and things; but, although the tend-
ency to individuation begins with life itself, it is completely
realized only in the higher animals.

622. In plants, as also in some of the lower animals, individu-
ality is merged in community. No plant (except one reduced
to the simplicity of a single cell, of circumscribed growth, and
without organs) is an individual in the sense that a man or a
dog is. (16, 156.) The herb, shrub, and tree are neither
indivisible nor of definite limitation. Whether their successive
growths are to remain parts of the previous plant, or to be inde-
pendent plants, depends upon circumstances ; and there is no
known limit to budding propagation.

623. There is, however, a kind of social or corporate indi-
viduality in those animals, or communities (whichever we call
them) of the lower grade which are multiplied by buds or off-
shoots as well as by ova, and in which the offspring remains, or
may remain, organically connected with the stock. The poly-
pidom or polyparium commonly has a certain limitation and a
definite form; and certain polyps may become organs with
special functions subordinate to the common weal. This is
more largely true in the vegetable kingdom. So that for de-
scriptive purposes, and in a just although somewhat loose sense,
the herb, shrub, or tree is taken as an individual. But only
while it forms one connected body. Offshoots when separately
established are equally individuals in this sense.

624. What it is in plants which philosophically answers to the
individual in the higher animals is another question, to which
various answers have been given. Some insist that the whole
vegetative product of one seed makes one individual, whether
connected or separated (as may happen) into a million of plants.
But a common and less strained view restricts the individual
to such product only while organically united. Others (of
which Thouars at the beginning and Braun at the middle of the
present century are leading examples) take each axis or shoot
with its foliage to represent the individual, of which the leaves
and their homologues are organs, the branches being usually
implanted upon the parent axis as this is implanted in the soil,
but also equally capable of producing roots by which they may
make their own connection with the soil. Still others, on pre-

1 For the history of opinion upon and a full presentation of this topic,
see Alexander Braun’s Memoir (originally published in the Abhandl. Akad.
Wissenschaften zu Berlin, 1853), Das Individuum der Pflanze, &c., and a
translation by C. F. Stone in Amer. Jour. Sci. ser. 2, xix. xx. 1855.

THE PRINCIPLES OF CLASSIFICATION. 317

cisely similar grounds, carry the analysis a step farther, and
regard each phytomer (16) as the individual. Finally, some, in
view of their potentially independent life, take the cells, or units
of anatomical structure, to be the true individuals; and this
with sufficient reason as regards the simplest cryptogamous
plants. Upon the view here adopted, that plants do not rise
high enough in the scale of being to reach true individuality,
the question is not whether it is the cell, the phytomer, the shoot,
the tree, or the whole vegetative product of a seed which answers
to the animal individual, but only which is most analogous to it.
In our view, its analogue is the cell in the lowest grades of vege-
table life, the phytomer in the higher.t Bnt, in botanical de-
scription and classification, by the individual is meant the herb,
shrub, or tree, unless otherwise specified.

625. Species in biological natural history is a chain or series
of organisms of which the links or component individuals are
parent and offspring. Objectively, a species is the totality ox
beings which have come from one stock, in virtue of that mosé
general fact that likeness is transmitted from parent to progeny.
Among the many deiinitions, that of A. L. Jussieu is one of the
briefest and best, since it expresses the fundamental conception
of a species, 7. e. the perennial succession of similar individuals
perpetuated by generation.

626. The two elements of species are: 1, community of origin ;
and, 2, similarity of the component individuals. But the degree
of similarity is variable, and the fact of genetic relationslip can
seldom be established by observation or historical evidence. — It
is from the likeness that the naturalist ordinarily decides that
such and such individuals belong to one species. Still the like-
ness is a consequence of the genetic relationship; so that the
latter is the real foundation of species.

1 For just as successive branches are repetitions and progeny of the
parent branch or stem, the phytomers of the branch are repetitions and
progeny each of the preceding one, so forming a series of vegetative
generations ; and the whole tree might almost as well represent the individ-
ual as one of its branches. The phytomer, as well as the branch, is capable
of completing itself by producing roots, but is itself indivisible except by
mutilation. Least tenable of all is the conception that the whole product
of a seed may be taken to represent the vegetable individual For then
individuals increased by buds and division are wholly unlimited both in ex-
tent and in duration, so far as observation can show, and a multitudinous
race, not only of the present and past, but perhaps in perpetuity, may con-
sist of a single individual. There are, indeed, theoretical reasons for infer-
ring that a bud-propagated race may not Jast so long as a seed-propagated
species ; but there is no proof of it. See Darwiniana, Art. xii.

318 TAXONOMY.

627. No two individuals are exactly alike; and offspring of
the same stock may differ (or in their progeny may come to differ)
strikingly in some particulars. So two or more forms which
would have been regarded as wholly distinct are sometimes
proved to be of one species by evidence of their common origin,
or more commonly are inferred to be so from the observation of
a series of intermediate forms which bridge over the differences.
Only observation can inform us how much difference is compat-
ible with a common origin. The general result of observation
is that plants and animals breed true from generation to genera-
tion within certain somewhat indeterminate limits of variation ;
that those individuals which resemble each other within such
limits interbreed freely, while those with wider differences do
not. Hence, on the one hand, the naturalist recognizes Varieties
or differences within the species, and on the other Genera and
other superior associations, indicative of remoter relationship of
the species themselves.

628. Varieties are forms of species marked by characters of
less fixity or importance than are the species themselves. They
may be of all grades of difference from the slightest to the most
notable: they abound in free nature, but assume particular
importance under domestication and cultivation; under which
variations are most prone to originate, and desirable ones are
preserved, led on to further development, and relatively fixed.

629. If two seeds from the same pod are sown in different
soils, and submitted to different conditions as respects heat, light,
and moisture, the plants that spring from them will show marks
of this different treatment in their appearance. Such differences
are continually arising in the natural course of things, and to
produce and increase them artificially is one of the objects of
cultivation. Striking as they often are (especially in annuals
and biennials), they are of small scientific consequence. When
spontaneous they are transient, the plant either outlasting the
modifying cause or else succumbing to its continued and graver
operation. But, in the more marked varieties which alone de-
serve the name, the cause is occult and constitutional; the
deviation occurs we know not why, and continues throughout
the existence and growth of the herb, shrub, or tree, and con-
sequently through all that proceeds from it by propagation from
buds, as by offsets, layers, cuttings, grafts, &e.

630. Some varieties of cultivation originate in comparatively
slight deviations from the type, and are led on to greater differ-
ences by strict selection of the most marked individuals to
breed from. Most appear as it were full-fledged. except as to

THE PRINCIPLES OF CLASSIFICATION. 319

luxuriance or development, more or less under the control of
conditions, their origin being wholly unaccountable. They arise
in the seed-bed, or sometimes from buds, which as the gardeners
say ‘‘ sport.”1 That is, some seedlings, or some shoots, are
unlike the rest in certain particulars.”

631. Most varieties originate in the seed, and therefore the
foundation for them, whatever it may be, is laid in sexual repro-
duction. But Bud-variation, or the ** 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
occur in corals, hydras, and other compound animals propagated
by budding. Once originated, these varieties mostly persist,
like seedling varieties, through all the generations of budding
growth, but are not transmitted to the seed.

632. Upon the general principle that progeny inherits or tends
to inherit the whole character of the parent, all varieties must
have a tendency to be reproduced by seed. But the inheritance
of the new features of the immediate parent will commonly be
overborne by atavism, z.e. the tendency to inherit from grand-
parents, great-grand-parents, &c. Atavisin, acting through a long
line of ancestry, is generally more powerful than the heredity
of a single generation. But when the offspring does inherit the
peculiarities of the immediate parent, or a part of them, its off-
spring has a redoubled tendency to do the same, and the next
generation still more; for the tendency to be like parent, grand-
parent, and great-grand-parent now all conspire to this result
and overpower the influence of remoter ancestry. Close-breed-
ing (398) is requisite to this result. In the natural wild state,
varieties — many and conspicuous as they often arc— must be
much repressed by the prevalent cross-fertilization which takes
place among tke individuals of almost all species. Cultivators and
breeders in fixing varieties are careful to secure close breeding
as far as this is possible. This has fixed the particular sorts of
Indian Corn, Rye, Cabbage, Lettuce, Radishes, Peas, &c., and

1 Both the technical English term. Sport, and its Latin equivalent, Lusus,
are sometimes used for bud-yariation only, yet as commonly for seedling
variation also.

2 Darwin assumes that variation is of itself indefinite or vague, tending
in no particular direction, but that direction is wholly given by the elimina-
tion in the struggle for life of all but the fittest for the conditions. But
what we observe in the sced-bed does not suggest this view. Negeli, Braun,
and myself incline to the opinion that each plant has an inherent tendency
to variation in certain general directions.

3 A list of known bud-varieties is given in Darwin’s Variation of Animals
and Plants under Domestication, Chapter xi.

320 TAXONOMY.

indeed of nearly all our varieties of cultivated annual and biennial
esculent plants, as well as of several perennials, many of which
have been fixed through centuries of domestication, while others
are of recent establishment. What is now taking place with
the Peach in this country may convince us that heritable varieties
may be developed in trees as well as in herbs, and in the same
manner; and that the reason why most races are annuals or
biennials is because these can be perpetuated in no other way,
and because the desired result is obtainable in fewer years than
in shrubs or trees. Varieties of this fixity of character are called

633. Races (Lat. Proles). A race, in this technical sense of
the term, is a variety which is perpetuated with considerable
certainty by sexual propagation. This distinction of varieties
pertains chiefly to botany. In the animal kingdom all permanent
varieties must be races. So are all indigenous varieties of
plants.’ In most of these, the position of species and variety is
more or less arbitrary or accidental, and capable of interchange.
What is called the species may be only a commoner or better-
known form, or the one first recognized and named by botanists ;
whence the other forms as they come to be recognized are made
to rank in the books as varieties. Instead of one varying from
the other, all the forms have probably varied ages ago from a
common type.

634. These varieties of the highest order and most marked
characteristics, being perpetuable by seed, have the principal
attributes of species. They are a kind of subordinate derivative
species. Hence they are sometimes called Subspecies. We
judge them not to be so many species, either because in the case
of cultivated races we know something of their origin or history,
and more of the grave changes which long domestication may
bring to pass; or because the forms, however stable, differ
among themselves less than recognized species generally do; or
because very striking differences in the extremes are connected
by intermediate forms. And our conclusions, it must be under-
stood, ‘‘are not facts, but judgments, and largely fallible judg-
ments.”? For while some varieties appear strikingly different,
some species are very much alike.*

! The Horseradish and a few other plants of spontaneous growth, which
through long dependence on bud-propagation seem to have lost the power
of setting seed, can hardly be called varieties.

2 Darwiniana, 35.

8 Wherefore, since we hardly need the term race in the restricted sense
of seed-propagated variety, it is sometimes convenient to use it in the man-
ner proposed by Bentham (Anniversary Address to the Linnean Society,

THE PRINCIPLES OF CLASSIFICATION. $21

635. One distinction between varieties and species is note-
worthy and important, even if it may not serve as a criterion.
The individuals of different varieties in plants interbreed as freely
as do those of the same variety and are equally prolific. Their
union produces

636. Cross-breeds.’ In nature, cross-breeding doubtless re-
presses variation or prevents the segregation of varicties into
what would be ranked as species. In cultivation and domesti-
cation, it is turned to important account in producing intermediate
new varieties (cross-breeds) variously combining the different
excellencies of two parent individuals or two varieties. Thus
the great number of forms produced by variation (especially as
to flowers and fruits) have been further diversified, and selected
forms improved for special uses by judicious combination.

637. In general, the individuals of distinct species do not
interbreed, although many are capable of it. There is great
diversity in this regard among plants, some (such as Willows,
Verbascums, and Verbenas) interbreeding freely and reciprocally ;
some interbreeding in one direction, but not reciprocally ; others,
even when very similar, refusing to unite. But, on the whole,
there seems to be few nearly related species in which the pollen
of the one cannot be made to act upon the ovules of the other
by persistent and proper management. Such crossing is an
important resource in horticulture. Crossing of species, when
successful, produces

658. Hybrids. In these, the characteristics of the two species
are combined, sometimes in equal proportions, sometimes with
great preponderance of one or the other parent; and there is
often a difference in the result in reciprocal fertilizations. Hy-
brids do not play a very prominent part in nature, apart from
cultivation, although the limits of some species may be obscured
by them, possibly of more than is generally supposed. In the
animal kingdom, all the most familiar hybrids are sterile: in the
vegetable kingdom, a majority may have a certain but very low
degree of fertility ; but this is also the case in many unions

May, 1869, 5) as the common designation of any group or collection of indi-
viduals whose characters are continued through successive generations,
whether it be permanent variety, subspecies, species, or group consisting
of very similar species, the term not implying any decision of this question.
If this use of the term race prevails, Subspecies will probably take its place
as the designation of the highest grade of variety. The objection to this is
that the subspecific and specific names would be more liable to be confused.
1 Half-breed is a common equivalent term in the animal kingdom: Latin,
Mistus or Miztus ; French, Métis.
2]

322, TAXONOMY.

within the species, and especially in the application of the pollen
to the stigma of the same blossom. Commonly the sterility of
hybrids is owing to the impotence of the stamens, which perfect
no pollen; and most such hybrids may be fertilized by the pollen
of the one or the other parent. ‘Then the offspring either in
the first or second generation reverts to the fertilizing species.
Moreover, certain hybrids, such as those of Datura, which are
fully fertile per se, divide in the offspring, partly in the first gen-
eration, and completely in two or three succeeding generations,
into the two component species, even when close-fertilized.1 (In
part this may come from adventive embryo-formation, 533.)

639. There appears, therefore, to be a real ground in nature
for species, notwithstanding the difficulty and even impossibility
in many cases of defining and limiting them.

640. Species is taken as the unit in zoological and botanical
classification. Important as varieties are in some respects,
especially under domestication and cultivation, they figure in
scientific arrangement only as fractions of species. Species
are the true subjects of classification. The aim of systematic
natural history is to express their relationship to each other.

641. The whole ground in nature for the classification of spe-
cies is the obvious fact that species resemble or differ from each
other unequally and in extremely various degrees. If this were
not so, if related species differed one from another by a constant
quantity, so that, when arranged according to their resemblances,
the first differed from the second about as much as the second
from the third, and the third from the fourth, and so on, — or if
the species blended as do the colors of the rainbow, — then, with
all the diversity in the vegetable kingdom there actually is, there
could be no natural foundation for their classification. The mul-
titude of species would render it necessary to classify them, but
the classification would be wholly artificial and arbitrary. The
actual constitution of the vegetable kingdom, however, as ap-
pears from observation, is that some species resemble each other
very closely indeed, others differ as widely as possible, and be-
tween these the most numerous and the most various grades of

! According to Naudin in Comptes Rendus, xlix. 1859, & lv. 1862. See
also Naudin’s memoir on hybridity in plants in Ann. Sci. Nat. ser. 4, xix.
1863, pp. 180-205, & in Mem. Acad. Sci. . . . For the literature on vegetable
hybrids, see Keelreuter, Nachricht, &c., 1761, and Appendices, 1763-1765;
Herbert, on Amaryllidacee, 1837 ; C. F. Gertner, Versuche und Beobachtun-
gen ueber die Bastarderzeugung in Pflanzenreich, 1849; Wichura, Die
Bastardbefruchtung im Pflanzenreich, erliutertert an den Bastarden der
Weiden ; and the memoir of Naudin referred to.

THE PRINCIPLES OF CLASSIFICATION. 323

resemblance or difference are presented, but always with a mani-
fest tendency to compose groups or associations of resembling
species, — groups the more numerous and apparently the less
definite in proportion to the number and the nearness of the
points of resemblance. ‘These various associations the naturalist
endeavors to express, as far as is necessary or practicable, 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. Linnzeus
and the naturalists of his day used names for only three grades
of association, or groups superior to species, viz. the Genus, the
Order, and the Class ; and these are still the principal members
of classification.

642. Genera (plural of Genus) are the more particular or
special groups of related species. They are groups of species
which are much alike in all or most respects, — which are con-
structed, so to say, upon the same particular model, with only
circumstantial differences in the details. They are not neces-
sarily nor generally the lowest definable groups of species, but
are the lowest most clearly definable groups which the botanist
recognizes and accounts worthy to bear the generic name; for
the name of the genus with that of the species added to it is the
scientific appellation of the plant or animal. _Constituted as the
vegetable and animal kingdoms are, the recognition of genera, or
groups of kindred species, is as natural an operation of the mind
as is the conception of species from the association of like indi-
viduals. This is because many genera are so strongly marked,
at least so far as ordinary observation extends. Every one
knows the Rose genus, composed of the various species of Roses
and Sweetbriers; the Bramble genus, comprising Raspberries,
Blackberries, &c., is popularly distinguished to a certain extent ;
the Oak genus is distinguished from the Chestnut and the Beech
genus; each is a group of species whose mutual resemblance is
greater than that of any one of them to any other plants. The
number of species in such a group is immaterial, and in fact is
very diverse. A genus may be represented by a single known
species, when its peculiarities are equivalent in degree to those
which characterize other genera. This case often occurs; al-
though, if this were universally so, genus and species would be
equivalent terms. If only one species of Oak were known, the
Oak genus would have been as explicitly discerned as it is now
that the species amount to three hundred; and better defined,
for now there are forms quite intermediate between Oak and

324 TAXONOMY.

Chestnut. Familiar illustrations of genera in the animal king-
dom are furnished by the Cat kind, to which belong the domestic
Cat, the Catamount, the Panther, the Lion, the Tiger, the Leop-
ard, &c.; and by the Dog kind, which includes with the Dog
the different species of Foxes and Wolves, the Jackal, &e. The
languages of the most barbarous as well as of civilized people
everywhere show that they have recognized such groups. Natu-
ralists merely give to them a greater degree of precision, and
indicate what the points of agreement are.

643. If most genera were as conspicuously marked as those
from which these illustrations are taken, genus would be as defi-
nitely grounded in nature as species. But popularly recognized
genera, rightly based, are comparatively few. Popular nomen-
clature, embodying the common ideas of people, merely shows
that generic groups are recognizable in a considerable number
of cases, but not that the whole vegetable or the whole animal
kingdom is divisible into a definite number of such groups of
equally or somewhat equally related species. The naturalist
discerns the ground of genera in characters which the casual and
ordinary observer overlooks; and, taking the idea of genera
from the numerous well-marked instances as the norm, applies
it as well as possible to the less obvious or less natural cases, and
groups all known species under genera. Resemblances among
the species when rightly grouped into genera, though real, are
often so unequal in degree, that certain species may be about as
nearly related to neighboring genera. So that the recognition of
genera even more than of species is a matter of judgment, and
even of conventional agreement as to how and where a certain
genus shall be limited, and what particular association of species
shall hold the position of genus. All the species of a genus must
accord in every important structure; but extended observation
only can settle the question as to what are important and what
are incidental characters. For example, the pinnatifid or sinu-
ate leaf might have been thought as essential to the Oak genus
as the acorn-cup; but many Oaks are now known with entire
leaves, resembling those of Willow or Laurel. An open acorn-
cup beset with imbricated scales is a character common to all
European and American Oaks; but in numerous Asiatic species
the cup bears concentric or spiral lamellze instead, and in others
the cup takes the form of a naked and closed sac. Maples have
palmately-veined and lobed leaves; but one species has undi-
vided and pinnately-veined leaves. The Apple and the Pear
under one view are of the same genus, under another they rep-
resent different genera.

THE PRINCIPLES OF CLASSIFICATION. Sy Ay

644. The genus must be based on close relationship of species,
but not necessarily on the closest. Raspberries differ from
Blackberries, but must be ranked in the same genus; and so of
Plums and Cherries. For the groups which are to bear the
generic name must be as distinct and definite as possible.

645. Orders are to genera what genera are to species. They
are groups of a higher rank and wider comprehension, expressive
of more general resemblances, or, in other language, of remoter
relationship. As all species must be ranked in genera, so all
genera must be ranked in orders. Famity in botany is synony-
mous with order: at least natural orders and families (however
distinguished in zoology) have always in botany been inter-
changeable terms, and will probably so continue.!

646. As examples of orders in the vegetable kingdom take
the Oak family, composed of Oaks, Chestnuts, Beeches, &e. ;
the Pine family, of Pines, Spruces, Larches, Cedars, Araucaria,
Cypresses, and their allies; the Rose family, in which Brambles,
Strawberries, Plums and Cherries, Apples and Pears are asso-
ciated with the Rose in one somewhat multifarious order.

647. Classes are to orders what these are to genera. They
express still more comprehensive relations of species ; each class
embracing all those species which are framed upon the same
broad plan of structure, however differently that plan may be
carried out in particulars.

648. Kingdom must be added, to represent the highest gener-
alization. All subjects of biological classification belong either
to the vegetable or animal kingdom. Mineralogy, Chemistry,
&¢c., may use the same terms (genus, species, &c.) in an analo-
gous way; but the classification of substances rests on other
foundations than that of beings.

649. The sequence of groups, rising from particular to univer-
sal, is Species, Genus, Order, Class, Kingdom; or, in descending
from the universal to the particular,

Kinepom,
Crass,
ORDER,
GENUS,
SPECIEs.

1 Order is the older term, and that which associates best with the technical
Latin names. Family is a happy term, which associates itself well with
English names. But its use is attended with this incongruity, that the tribe
(653) in natural history classification is subordinate to the family. In
zoology, order is distinguished from family as the next higher grade.

326 TAXONOMY.

650. This is the common framework of natural history classi-
fication. All plants and all animals belong to some species ;
every species to some genus; every genus to some order or
family ; every order to some class; every class to one or the
other kingdom.t But this framework, although all that is re-
quisite in some parts of natural history, does not express all the
observable gradations of relationship among species. And even
gradations below species have sometimes to be classified. The
series is capable of extension; and extension is often requisite
on account of the large number of objects to be arranged, and
the various degrees of relationship which may come into view.

651. This is effected by the intercalation of intermediate
grades, to be introduced into the system only when there is
occasion for them. And in botany one or more grades superior
to the classes are needful; for first and foremost is the great
division of all plants into a higher and a lower Series? (or sub-
kingdom), the Phzenogamous and the Cryptogamous.

652. The grades intercalated into the long-established sequence
of Class, Order, Genus, and Species, with new names, are mainly
two, Tribe and Cohort.

653. Tribe has been for a generation or two thoroughly estab-
lished in both kingdoms, as a grade inferior to order and supe-
rior to genus. In botanical classification, much use is made of
this grade, genera being grouped into tribes.

654. Cohort (Lat. Cohors) is of more recent introduction, at
least in Botany, but is becoming established for a grade next
above that of order. Orders are grouped into cohorts. Lindley
hit upon a good English name for this grade, that of Alliance.
But this word has no available Latin equivalent; while cohort
takes equally well a Latin or an English form.

655. Finally, each grade is capable of being doubled by the
recognition of one like it and immediately subordinate to it, and
with designation directly expressive of the subordination. For

1 Not recognizing Heckel’s third kingdom of Protista, consisting of those
lowest forms of being from which the animal and vegetable kingdoms
emerge.

2 Answering to the French Embranchement in zoology. For this it is pro-
posed to use the word Dyirision (Divisio): see Laws of Botanical Nomencla-
ture adopted by the International Botanical Congress held at Paris in
August, 1867 ; together with a Historical Introduction anda Commentary, by
Alph. DeCandolle, — English translation, London, 1868 ; the original French
edition, Paris, 1867. Perhaps no better name can be found; but the elder
DeCandolle brought Divisio into common use for a grade subordinate to
tribe. Endlicher employed the term Regio. We have used Series, and
much prefer it.

THE PRINCIPLES OF CLASSIFICATION. 327

example, if Dicotyledones and Monocotyledones be the two
classes of Phznogamia, the former (and only the former) is
divided upon very important characters into two branches, of far
higher rank than the cohorts, viz. the Angiosperme and the
Gymnosperme, which take the name of Sunctasses. Orders,
especially the more comprehensive ones, often comprise two or
more groups so distinct that it may fairly be a question whether
they are not of ordinal rank: such take the name of SuBorDERs.
Tribes in like manner may comprise groups of similar relative
value: these are Suprrises. Genera may comprise sections of
species which might almost as well rank as genera themselves :
to mark their importance and pretension (which may come to
be allowed), they are termed SusGenera. Finally, forms which
are ranked as varieties, but which may establish a claim to be
distinct species, are sometimes termed Scupspecres. Even what
we regard as a variety may comprise more or less divergent
forms, to be distinguished as SUBVARIETIES.

656. Some of the larger and most diversified orders, tribes,
genera, or species may require all these analytical appliances,
and even more, for their complete elucidation; while others,
comparatively homogeneous, offer no ground for them. But
when these grades, or some of them, come into use, they are
always in the following sequence : —

KINGDOM,
Series or Division, or Sub-kingdom,
Grass:
Subclass,
Cohort,
OrD_ER or Family,
Suborder,
TRIBE,
Subtribe,
GENUS,
Subgenus,
Section,
Subsection,
SPECIES,
Subspecies or Race,
Variety,
Subvariety.

657. Nature and Meaning of Affinity. These grades, the higher
including the lower, denote degrees of likeness or difference.
Plants belonging respectively to the two great series or primary
divisions may accord only in the most general respects, in that
which makes them plants rather than animals. Plants of the
same variety are generally as much alike as if they were of the

328 TAXONOMY.

same immediate parentage. All plants of the same species are
so much alike that they are inferred to have descended from
a common stock, and their differences, however graye, are sup-
posed to have arisen from subsequent variation, and the more
marked differences to have become fixed through heredity. This
is included in the idea of species. Descent from a common
origin explains the likeness, and is the only explanation of it.

658. But what is the explanation of the likeness between the
species themselves? As respects nearly related species, the answer
is clear. Except for practical purposes and in an arbitrary
way, no certain and unfailing distinction can be drawn between
varieties of the highest grade and species of closest resemblance.
It cannot reasonably be doubted that they are of similar origi-
nation. Then there are all gradations between very closely and
less closely related species of the same genus of plants.

659. The Theory of Descent, that is, of the diversification of the
species of a genus through variation in the lapse of time, affords
the only natural explanation of their likeness which has yet been
conceived. The alternative supposition, that all the existing
species and forms were originally created as they are, and have
come down essentially unchanged from the beginning, offers no
explanation of the likeness, and even assumes that there is no
scientific explanation of it. The hypothesis that the species of
a genus have become what they are by diversification through
variation is a very old one in botany, and has from time to time
been put forward. But until recently it has had little influence
upon the science, because no clear idea had been formed of any
natural process which might lead to such result. Doubtless, if
variation, such as botanists have to recognize within the species,
be assumed as equally or even more operative through long ante-
rior periods, this would account for the diversification of an
original species of a genus into several or many forms as differ-
ent as those which we recognize as species. But this would not
account for the limitation of species, which is the usual (but
not universal) characteristic, and is an essential part of the
idea of species. Just this is accounted for by

660. Natural Selection. This now familiar term, proposed by
Charles Darwin, was suggested by the operations of breeders in
the development and fixation of races for man’s use or fancy ;—
in animals by breeding from selected parents, and selecting for
breeding in each generation those individuals only in which the
desired points are apparent and predominant; in the seed-bed
by rigidly destroying all plants which do not show some desirable
variation, breeding in and in from these, with strict selection of

THE PRINCIPLES OF CLASSIFICATION. 3829

the most variant form in the particular line or lines, until it be-
comes fixed by heredity and as different from the primal stock
as the conditions of the case allow. In nature, the analogous
selection, through innumerable generations, of the exceedingly
small percentage of individuals (as ova or seeds) which ordi-
narily are to survive and propagate, is made by competition for
food or room, the attacks of animals, the vicissitudes of climate,
and in fine by all the manifold conditions to which they are
exposed. In the Struggle for Life to which they are thus inevi-
tably exposed, only the individuals best adapted to the circum-
stances can survive to maturity and propagate their like. This
Survival of the Fittest, metaphorically expressed by the phrase
natural selection, is in fact the destruction of all weaker com-
petitors, or of all which, however they might be favored by other
conditions, are not the most favored under the actual circum-
stances. But seedlings varying, some in one direction, some in
another, are thereby adapted to different conditions, some to one
kind of soil or exposure, some to another, thus lessening the com-
petition between the two most divergent forms, and favoring their
preservation and farther separation, while the intermediate forms
perish. Thus an ancestral type would become diversified into
races and species. Earlier variation under terrestrial changes
and vicissitudes, prolonged and various in geological times since
the appearance of the main types of vegetation, and the attendant
extinctions, are held to account for genera, tribes, orders, &c.,
and to explain their actual affinities. Affinity under this view
is consanguinity ; and classification, so far as it is natural, ex-
presses real relationship. Classes, Orders, Tribes, &c., are the
earlier or main and successful branches of the genealogical tree,
genera are later branches, species the latest definitely developed
ramifications, varieties the developing buds.!

661. Except as to those changes in size, luxuriance, or depau-
peration and the like, in which plants, especially seedlings,
respond promptly to external influences, as to heat or cold,

1 For the inception of this theory of descent in the form which has within
the last twenty years profoundly affected natural history, and developed a
copious literature, see a short paper On the Variation of Organic Beings in
a State of Nature; On the Natural Means of Selection; and On the Com-
parison of Domestic Races and True Species, by Charles Darwin, also On
the Tendency of Varieties to depart indefinitely from the Original Type, by
Alfred Russell Wallace, both read to the Linnean Society, July 1, 1858, and
published in its Journal of the Proceedings, iii. (Zoology) 45-62. For the
development of the doctrine, see Darwin’s “ Origin of Species by Means of
Natural Selection,’ “The Variation of Animals and Plants under Domes-
tication,” and various other works; Wallace’s “Geographical Distribution
of Animals,” &c. For some expositions, see Gray’s “ Darwiniana.”

350 TAXONOMY.

moisture or dryness (which are transient and comparatively
unimportant), variation, or the unlikeness of progeny to parent,
is occult and inexplicable. If sometimes called out by the
external conditions, it is by way of internal response to them.
In Darwin’s conception, variation of itself does not tend in any
one particular direction: he appears to attribute all adaptation
to the sorting which results from the struggle for existence and
the survival of the fittest. We have supposed, and Negeli takes
a similar view,’ that each plant has an internal tendency or pre-
disposition to vary in some directions rather than others; from
which, under natural selection, the actual differentiations and
adaptations have proceeded. Under this assumption, and taken
as a working hypothesis, the doctrine of the derivation of species
serves well for the co-ordination of all the facts in botany, and
affords a probable and reasonable answer to a long series of
questions which without it are totally unanswerable. It is sup.
ported by vegetable paleontology, which assures us that the
plants of the later geological periods are the ancestors of the
actual flora of the world. In accordance with it we may explain,
in a good degree, the present distribution of species and other
groups over the world. It rationally connects the order of the
appearance of vegetable types in time with the grades of differ-
entiation and complexity, both proceeding from the simpler, or
lower and more general, to the higher and more differentiated
or special; it explains by inheritance the existence of function-
less parts; throws light upon the anomalies of parasitic plants in
their various gradations, upon the assumption of the most various
functions by morphologically identical organs, and indeed illumi-
nates the whole field of morphology with which this volume has
been occupied. It follows that species are not ‘+ simple curiosities
of nature,” to be catalogued and described merely, but that they
have a history, the records of which are impressed upon their
structure as well as traceable in their geographical and paleon-
tological distribution. This view, moreover, explains the re-
markable fact that the characters in which the affinities of
plants are mainly discerned (and which therefore serve best
for orders, tribes, and other principal groups) are commonly such
as are evidently of small if any importance to the plant’s well-
being, and that they run like threads through a series of species
of the greatest diversity in habit, mode of life, and particular
adaptations to conditions.’

1 Entstehung und Begriff der naturhistorischen Art. Zweite Auflage, 1865.
2 This is a corollary of natural selection, which can take effect only
upon useful characters, 7. ¢. upon structures which play some active part

BOTANICAL CLASSIFICATION. 331

662. The fixity of species under this view is not absolute and
universal, but relative. Not, however, that specific changes are
necessitated in virtue of any fixed or all-controlling natural law.
Some of the lowest forms have existed essentially unchanged
through immense geological periods down to the present time ;
some species even of trees are apparently unchanged in the lapse
of time and change of conditions between the later tertiary period
and our own day, during which most others have undergone
speciiic modification. Such modifications are too slow to effect
in any wise the stability and practical application of botanical
classification.

Section II. BorantcaLt CLASssIFICATION.

663. Natural and Artificial Classifications may be distinguished.
A natural classification in botany aims to arrange all known plants
into groups in a series of grades according to their resemblances,
and their degrees of resemblance, in all respects, so that each
species, genus, tribe, order, &c., shall stand next to those which
it most resembles in all respects, or rather in the whole plan of
structure. For two plants may be very much alike in external
appearance, yet very diiferent in their principal structure. Arti-
ficial classifications single out one or more points of resemblance
or difference and arrange by those, without reference to other
considerations, convenience and facility being the controlling
principles. The alphabetical arrangement of words in a dic-
tionary, and the sexual system in botany by Linnzeus (or rather
a part of it), in which plants are arranged in classes upon the
number of their stamens, and in orders upon the number of
pistils, —are examples of artificial classification. The arrange-
ment of the words of a language under their roots, and with the
derivative under the more primitive forms, would answer to a
naturai classification.

in the life of the plant, and which therefore undergo modification under
changing conditions. Unessential structures accordingly are left unaltered
or are only incidentally modified. And so these biologically unessential
points of structure, persisting through all adaptive changes, are the clews
to relationship. Thus, Rubiacee are known by insignificant stipules, Ano-
nace by ruminated albumen, Rhamnacee by a valvate calyx and stamens
before the petals, &e. Paradoxicat as it may seem, it is not although, but
because, they are of small biological importance that they are of high clas-
sificatory (7. e. of genealogical) value.

On considerations like these, characters are divided into adaptive or bio-
logical on the one hand, and genealogical or genetic on the other. The saga-
cious naturalist seizes upon the latter for orders and the like; while the
former are prominent in genera, &e. ii

332 TAXONOMY.

664. No artificial classification of plants could fail to be
natural in some portions and some respects; because plants
whick agree in any point of structure likely to be used for the
purpose will commonly agree in other and perhaps more impor-
tant characters. On the other hand, no natural classification can
dispense with artificial helps; nor can it express in lineal order,
or in any other way, all the various relationships of plants, even
if these were fully determined and rightly subordinated. Natu-
ralists now endeavor to make classification as natural as possible ;
that is, to base it in every grade upon real relationships. What
real relationships are, and how to express them in a general
system and throughout its parts, has been the task of the leaders
in botany from the beginning of the science until now; and the
work is by no means completed.

665. Linnzus was perhaps the first botanist to distinguish
clearly between a natural and an artificial classification. He
labored ineffectually upon a natural classification of the genera
of plants into orders ; and he devised an effective artificial classi-
fication, which became so popular that it practically superseded
all others for more than half a century, and has left a permanent
impression upon the science. The last generation of botanists
who were trained under it has not quite passed away.

666. Ante-Linnean Classification. Linnzeus, in his Philosophia
Botanica, divided systematists into heterodox and orthodox:
the former, those who classify plants by their roots, herbage, time
of flowering, place of growth, medical and economical uses, and
the like ; the latter, by the organs of fructification. It is remark-
able that all the orthodox or scientific classifications anterior to
Linneus made a primary division of the vegetable kingdom into
Trees and Herbs, referring the larger shrubs to the former and
the under-shrubs to the latter,—an arrangement which began
with Theophrastus and was continued by Ray and Tournefort.

667. The three most important names in botanical taxonomy
anterior to Linnzeus are those of Cesalpini, Ray, and Tournefort.
Scientific botany commenced with the former, in Italy, in the
latter half of the sixteenth century. He first used the embryo and
its cotyledons in classification. distinguished differences in the in-
sertion of floral parts, and. indeed (excepting the primary division
into trees and herbs). founded all principal characters upon the
organs of fructification, especially upon the fruit and seed.
Conrad Gesner of Zurich had somewhat earlier recognized this
principle, but Cesalpini first applied it.

668. A century later (1690-99) this principle was carried into
practice by Rivinus (a name latinized from Bachmann), of

BOTANICAL CLASSIFICATION. 333

Leipsic, in a wholly artificial classification founded on the corolla.
His contemporary in England, Robert Morison, somewhat earlier
began the publication of his great work, the Universal History
of Plants. In this was first attempted a grouping of plants into
what are now called natural orders ; and these were defined, some-
what loosely, some by their fruit, inflorescence, and flowers, others
by their stems, the nature of their juice, &c. But the two great
systematists of the time, who together laid the foundations of
modern scientific botany, were John Ray in England and Joseph
Pitton de Tournefort in France.

669. 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 subdivided, by characters taken from the organs of fructi-
fication, into what we should cail natural orders or families, but
which he unfortunately called genera. He noted the coincidence
of nerved leaves with the monocotyledonous embryo, although
he did not notice that his first division of arborescent plants was
monocotyledonous ; and he had a clear apprehension of genera.

670. Tournefort’s method was published in French in the
year 1694, in Latin in 1700. It is more definite but more arti-
ficial than that of Ray, being founded like that of Rivinus almost
wholly upon modifications of the corolla, and it overlooked the dis-
tinction between monocotyledonous and dicotyledonous embryos.
Its great merit is that here genera, as we now understand them,
are first established and defined, and all the species then known
referred to them; so that Tournefort was justly said by Linnzus
to be the founder of genera. Ray may be said to have indicated
the primary classes, Jussieu (in the next century) to have estab-
lished natural orders, and Tournefort to have given to botany
the first Genera Plantarum.

671. Linnean Classification. Linnzeus, the great reformer of
hotany in the eighteenth century, thoroughly revised the principles
of classification, established genera and species upon a more scien-
tific basis, and, in designating species by a word instead of a
descriptive phrase, introduced binomial nomenclature. (704.)
He likewise established for-the stamens, and indeed for the
pistils also, their supreme importance in classification (probably
without knowledge of the clear suggestion to this effect made
by Burckhard in a letter to Leibnitz, printed in 1702) ; their
functions, so long overlooked, being now ascertained. He also

334 TAXONOMY.

drew a clear and practical distinction between natural and arti-
ficiai classifications (663), and deferring all endeavors to make
the former available, except for genera, he devised a practical
substitute for it, as a key to the genera, viz. his celebrated

672. Sexual System, or arrangement of the genera under arti-
ficiai classes and orders, founded upon the stamens and pistils.
Although now out of use, this artificial classification has been so
popular and influential, and has left so deep an impression upon
the science and especially upon the language of botany, that it
needs to be presented. The primary divisions are the classes,
twenty-four in number. But the 24th class, Cryptogamia, con-
sists of plants which have not stamens and pistils and conse-
quently no proper flowers, and is therefore the counterpart of the
remaining twenty-three classes, to which the corresponding name
of Phanerogamia or, in shorter form, Pheenogamia (Phanogamous
plants} has since been applied. These twenty-three classes are
characterized by certain modifications and associations of the
stamens, and have substantive names, of Greek derivation, ex-
pressive of their character. The first eleven comprise all plants
with perfect (7. e. hermaphrodite) flowers, and with a definite
number of equal and unconnected stamens. ‘They are distin-
guished by the absolute number of these organs, and are desig-
nated by names compounded of Greek numerals and the word
andria (from «r7je), which is used metaphorically for stamen, as
follows : —

Class 1. Monanprta includes all such plants with one stamen to the flower;
as in Hippuris.

Dranpria, those with two stamens, as in the Lilac.

TRIANDRIA, with three stamens, as in the Valerian and Iris.

TETRANDRIA, with four stamens, as in the Scabious.

PENTANDRIA, with five stamens, the most frequent case.

HEXANDRIA, with six stamens, as in the Lily Family, &e.

HeEpPTANDRIA, with seven stamens, as in Horsechestnut.

OcranbrRIA, with eight stamens, as in Evening Primrose and Fuchsia.

. ENNEANDRIA, with nine stamens, as in the Rhubarb.

. DEcANDRIA, with ten stamens, as in Rhododendron and Kalmia.

DopecaNnpDRIA, with twelve stamens, as in Asarum and the Migno-
nette; extended also to include those with from thirteen to nine-
teen stamens.

Fee, SCOR TE OO ON ES CONS

et

673. The two succeeding classes include plants with perfect
flowers having twenty or more unconnected stamens, which, in

12. IcosanpRIA, are inserted on the calyx (perigynous), as in the Rose
Family; and in

13. Potyanprta, on the receptacle (hypogynous), as in the Buttercup,
Anemone, &c.

BOTANICAL CLASSIFICATION. ae

674. Their essential characters are not indicated by their
names: the former merely denoting that the stamens are twenty
in number; the latter, that they are numerous. — The two fol-
lowing classes depend upon the relative length of the stamens,
namely : —

14. Dipynamrta, including those with two long and two short stamens,
as in the majority of flowers with bilabiate corolla.

15. TerTRApYNAMr1a, those with four long and two short stamens, as in
flowers with cruciferous corolla.

675. These names signify in the former that two stamens, and
in the latter that four stamens, are most powerful. — The four
succeeding are founded on the connection of the stamens, viz. : —

16. MonapELPuHIA (meaning a single fraternity), with the filaments
united in a single set, tube, or column, as in the Mallow.

17. DiapeLruia (two fraternities), with the filaments united in two
sets or parcels, as in Corydalis and in many Leguminose.

18. PotyapEeLpuia (many fraternities), with the filaments united in
more than two sets or parcels, as in Hypericum.

19. Syncenesia (from Greck words signifying to grow together),
with the anthers united in a ring or tube, as in the Sunflower
and all Composite.

676. The next class, as its name denotes, is founded on the
union of the stamens to the style : —

20. GyNANDRIA, with the stamens and styles consolidated, as in Cypri-
pedium and all the Orchis Family.

677. In the three following classes, the stamens and pistils
occupy separate blossoms : —

21. Monecia (one household) includes all plants where the stamens
and pistils are in separate flowers on the same individual; as in
the Oak and Chestnut.

22. Diacia (two households), where they occupy separate flowers on
different individuals ; as in the Willow, Poplar, Moonseed, &e.

23. PotyGamia, where the stamens and pistils are separate in some
flowers and associated in others, either on the same or two or
three different plants; as in most Maples. *

678. The remaining class is essentially flowerless ; or rather its
organs of reproduction are more or less analogous to, but not
homologous with, stamens and pistils. But, although Linnzeus
suspected a sexuality in Ferns, Mosses. Algze, &c., there was no
proof of it in his day. So he named the class, containing these,

.24, CryproGAmiA, 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 : —

TAXONOMY.

336

‘VINVOOLAANO
‘VINVDATIOY

*VIOWIC
*VIOIDNO IAL
*VINGNVNAY

‘VIS

UO NAG

‘VIHATIAVAITOd

VINATAOV ICT
‘VIHA THOVNO[L

“VINVNAGVULLAL,

“VIN VNACIC

‘VINGNVATOT
“VIMGNVSOOT
“VIMNAUNVOMCOCT
“VINANVOCUC
*VINGNVANNG
“VINANVLOO
“VINGNV LAT]
“VINGNVXGIT
“VINGNV ING
VINAINVULAT,
“VINGNVIWT,
“VIMAINVICT
“VINGNVNO|A

sTeNpIA
=IPUT JUSIAYIP O91} IO OM} IO ‘ours oy UL
‘poyurudes 8101130 ‘yoaytod S1OMOTF Oo) JO ONLOS
as SPEUPIATpUL JUdLOYIp UL
* S[UNPLATpUL ous ot[} UL

“saoyue aporyy Aq
8}08 omy UvT} O10 UL syuUOUTETYy toy Aq

S}OS OMY UL SPUOUULL TT Tot) Aq
408 OLSUIS UV UL S}UOMIE]Y Afolyy Aq

SUOTILYS JLOYS OMY PUB FuoyT AMoJ
* SUOTUUS JALOT[S OA PUB SUOL OMY

xA7TwO OT) 09 YUdAOYpY you foroNr 10 TZ
* * xA[VO O17) 07 queLOT|pY ‘OMOUL 10 0% A
. . . . . . . . . . 6b II 4
; ; pen Ot: gape OL "
. . . . . . . . . . . . . L “ce
. . . . . . . . . . . . . ¢ “e
. . . . . . . . . . . . . ¢ “
. (pan o* ik . pee . ps . . ¢ “
. . . . . . . . . . . . . 4 oe

Foe fe.” sy fee ere! oteram ee Or ib

I suoulNys

‘SUSSVIO NVWNNI'T

es

OUHAL

Sepia Al eer er oh Aly.

auou 10 ‘poyvoouod s[ystd puv suouinys o1}

* g19MOTT 0}BrEdoS UT

* Tys8td oY 07 JUaIOYpV suoUTRyS OT) }

PY suoy
jenboun jo

| pur ‘10190
youo IIA

* TOTO TPHVO YQEM poyoouT0

pojoouuooun

Tysuoy
wubo jo

‘TOMO
oles OY} UT
‘ssid punoy y40q
OL} WLOAy
OPV
SudUIeySs
oy

TO MUTA TVOLLCONAS

“qsoyTue ur
ststd puv
suotUeys

Suavy
SINVIG

BOTANICAL CLASSIFICATION. 337

680. The orders, in the first thirteen classes of the Linnean
artificial system, depend on the number of styles, or of the
stigmas when the styles are wanting; and are named by Greek
numerals prefixed to the word gynia, used metaphorically for
_pistil, as follows : —

Order 1. Monocynta, those with one style or sessile stigma to the flower.
2. Dicyni4, those with two styles or sessile stigmas.
3. TricyniA, those with three styles.

4. TerracyniA, those with four styles.
5. PentaGYNiA, those with five styles.
6. HexaGynta, those with six styles.
7. HeprTacynia, those with seven styles.
8. OcroeyniA, those with eight styles.
9. EnneacyntA, those with nine styles.
10. DecaGynta, those with ten styles.
11. Dopecacynta, those with eleven or twelve styles.
12. Potyeynta, those with more than twelve styles.

681. The orders of class 14, Didynamia, are only two and are
founded on the pericarp, namely : —

1. GyMNOSPERMIA, meaning seeds naked, the achenia-like fruits of a
4-parted pericarp having been taken for naked seeds.

2. ANGIOSPERMIA, With the seeds evidently in a sced-vessel or peri-
carp, 7. e. the pericarp undivided.

682. The 15th class, Tetradynamia, is also divided into
two orders. which are distinguished merely by the form of the
pod : —

1. Srxricutosa; the fruit a silicle (561), or short pod.
2. Siz1quosa; fruit a silique (561), or more or less elongated pod.

683. The orders of the 16th, 17th, 18th, 20th, 21st, and 22d
classes depend merely on the number of stamens ; that is, on the
characters of the first thirteen classes, whose names they likewise
bear: as Monanpria, with one stamen, Dianpria, with two
stamens ; and so on.

684. The orders of the 19th class, Syngenesia, are six, namely :

1. PoLyGAMIA £QUALIS, where the flowers are in heads (the so-called
compound flower), and all hermaphrodite.

2. POLYGAMIA SUPERFLUA, the same as the last, except that the rays,
or marginal flowers of the head, are pistillate only.

3. POLYGAMIA FRUSTRANEA, those with the marginal flowers neutral,
the others perfect.

4 PoLyGAaMiA NECESSARIA, where the marginal flowers are pistillate

_ and fertile, and the central staminate and sterile.

5. POLYGAMIA SEGREGATA, where each flower of the head [or glom-
erule] has its own proper involucre.

6. Monocamta, where solitary flowers (that is, not united into a head)
have united anthers, as in Lobelia.

338 TAXONOMY.

685. The 23d class, Polygamia, has three orders, two of them
founded on the characters of the two preceding classes and
bearing their names, and the third named upon the same prin-
ciple, namely : —

1. Mone@cra, where both separated and perfect flowers are found in
the same plant.
. Diacra, where they occupy two different plants.

. Tri@cra, where one individual bears the perfect, another the stami-
nate, and a third the pistillate flowers.

ww bo

686. The orders of the 24th class, Cryptogamia, the Flower-
less Plants, are so many natural orders, and are not definable
by a single character. They are : —

1. Frrices, the Ferns.

2. Musct, the Mosses.

3. ALG#, which, as left by Linnzus, comprised the Hepatice, Lichens,
&e., as well as the seaweeds.

4. Fune1, Mushrooms, &e.

687. In its day, this artificial system well fulfilled its purpose,
and was preferred to all others on the score of facility and defi-
niteness. Now no botanist would think of employing it, nor
would it be chosen for a key to genera, which was its only legiti-
mate use.

688. The Natural System was rightly appreciated by Linnzeus,
who pronounced it to be the first and last desideratum in syste-
matic botany; and he early attempted to collocate most known
genera under natural orders (e. g. Piperite, Pulme, Scitamine,
Orchidee, Amentacea, &c., sixty-seven in number, including his
four cryptogamic orders), but without definition or arrangement.
In his later years, he was unable to accomplish any thing more.
The difficult problem was taken up by Linnzeus’s contemporary
and correspondent, Bernard de Jussieu, who planted the botanic
garden at Trianon with plants grouped into natural orders, but
published nothing. His pupil, Adanson, who when a young
man lived for several years in Senegal, and who was as remark-
able for eccentricity as for erudition and ability, published in
1763, in his Familles des Plantes, the first complete system of
natural orders. But he seems to have taken little from his
teacher, and with all his genius to have contributed little to the
advancement of the natural system.

689. Antoine Laurent de Jussieu, nephew of Bernard, has
been called the founder of the natural system of botany, and to
him more than to any other one person this honor may be
ascribed. In his Genera Plantarum secundum Ordines Natu-
rales disposita, 1789, natural orders of plants, one hundred

Pe ee ee ee Cle eee ee ee ae

BOTANICAL CLASSIFICATION. 339

in number, were first established and defined by proper char-
acters, and nearly all known genera arranged under them. His
primary division of the Vegetable kingdom was into Acotyledones,
Monocotyledones, and Dies yledones, adopted from Ray, with a
change which was no improvement. For his Acotyledones, the
Cryptogamia of Linnzeus, 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 embryo altogether.
The Acotyledones forming his first class, Jussieu divided the
Monocotyledones into three classes upon single and artificial
characters, namely upon the insertion of the stamens, whether
hypogynous, perigynous, or epigynous ; and the Dicotyledones,
into eleven classes on similar characters, preceded by a division
into Apetale, Monopetale, Polypetale, and Diclines irregulares, i.e.
first upon the character of the perianth, then upon the insertion
of the stamens or in Monopetalee of the corolla. The following
is the scheme : —

Acotyledones. . . He See «ces CLASS OI:
Stamens hypogynous . . . . . If.
Monocotyledones perigynouses oe. 2. EE
epipynousia as os. <= live

Stamens epigyuouss e723) 0 127" V.

Apetalous . PeLioynousy 4 Gece 30 ee VAL

by pooynoust A ac oe, VE

Corolla hypogynous . . . . .VIII.

perigynous . Bore corel D.&

epigynous: anthers connate X.

epigynous: anthers separate XI.

(

Monopetalous
Dicotyledones . 4

(

Stamens epigynous. . .. . . XII.

Polypetalous . hypogynous . .. . XIII.
perigynousi:, |. 8.52, 4 UNLV.

Diclinous (also Apefalous) - . =. .- . . . . . 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 began with the latter, the pheenogamous or flowering plants,
and with those having typically complete flowers. On account
of its convenience and the greater facilities for studying the higher
plants, this order has been commonly followed ever since. His
primary division on anatomical structure, into Vascular and
Cellular plants. was a backward step, confusing a portion of the
lower series with the higher; and the duplicate names of Exo-
gene and Endogene, appended to Dicotyledones and Monoco-

340 TAXONOMY.

tyledoneze, as it now appears should have been omitted. The
giades of the Candollean system superior to the orders, in their
final form, are mainly these : —

pry. I. VASCULAR (more properly PHA2NOGAMOUS) PLANTS.

Crass I. DicoryLeponowus or EXOGENOUS.
Subclass I. THaLtamirLorous : petals /distinct) and stamens on the
torus, 7. e. free.
II. Catyciritorous: petals (distinct or coalescent) and
stamens adnate to the calyx.

III. CorotiirLorous: petals (mostly coalescent) not ad-
nate to calyx, bearing the stamens.

IV. Monocutamypeous: petals wanting.

Crass IL. MonocotyLeponous or ENpoGENous. (No subclasses.)

Diy. Il. CELLULAR (more properly CRYPTOGAMOUS) PLANTS.
Crass I. ASTHEOGAMOUS: with sexual apparatus, and
Vascular tissue. (Hquisetacew—Filices.)
Only cellular tissue. (Musci and Hepatice. )

Cuiass II. AmpuiGamous: destitute of sexual organs and of other than
cellular tissue. (Lichenes, Fungi, Alge.)

691. Cryptogamous plants of all orders are now known to be
provided with sexes; and the Jussixan divisions of the Dico-
tyledones into Apetale (including Diclines), Monopetale, and
Polypetale, are generally preferred to those of DeCandolle. Into
the present views of the classification of the Cryptogamia it is
unnecessary here to enter. Their general arrangement into
classes, &c., is not yet well settled, and the whole taxonomy
of the lower Cryptogams is at present in a state of transition.

692. John Lindley in successive attempts (between 1830 and
1845) variously modified, and in some few respects improved,
the Candollean arrangement. But, as neither his groupings of the
natural orders nor the new classes which he adopted have been
approved, his schemes need not be here presented. He must be
credited, however, with the first attempt to carry into effect a
suggestion made by Brown, that the orders should themselves
be disposed as far as possible into superior and strictly natural
groups. In Lindley’s first attempt, such groups of two grades
were proposed, the lower called nixus (tendencies), the higher
cohorts. In his later and largest work, The Vegetable King-
dom, these were reduced to one, and the name of alliance was
coined. But this word has no good Latin equivalent, and the
term cohort (coors) is preferred.

BOTANICAL CLASSIFICATION. 341

693. Robert Brown, next to Jussieu, did more than any other
botanist for the proper establishment and correct characterization
of natural orders. Having in the year 1827 published his dis-
covery of the gymnospermy of Coniferse and Cycadacez, it
was in Lindley’s works that this was first turned to proper
systematic account by dividing the class of Dicotyledones into
two subclasses, the Angiosperme and the Gymnosperme. The
latter has been elevated by the vegetable paleontologists to the
rank of a class.

694. Stephen Ladislaus Endlicher, of Vienna, a contempo-
rary of Lindley, of less botanical genius, but of great erudition
and aptness for classification, brought out his complete Genera
Plantarum secunduin Ordines Naturales disposita, between the
years 1836 and 1840. This elaborate work follows that of its
predecessor, Jussieu, in beginning with the lower series of plants
and ending with the higher. Its primary division is into two
regions: 1. Thallophyta, plants without proper axis of growth
(developing upward as stem and downward as root), no other
tissue than parenchyma, and (as was thought) no proper sexes.
This answers to the lower or Amphigamous Cellular plants of
DeCandolle. 2. Cormophyta, plants with an axis (stem and root),
with foliage, &c. The Cormophyta, or plants of the higher
region, Endlicher divided into three great sections: 1. Acro-
brya, answering to the higher A%theogamous Cryptogamia of
DeCandolle, with which was wrongly associated a group of root-
parasitic flowering plants (the Rhizanthez) which were fancied
to bear spores instead of embryo in their seed; 2. Amphibrya,
which answer to Monocotyledones ; and, 3. Aeramphibrya, which
answer to Dicotyledons. These last contain five cohorts: 1.
Gymnospermee ; 2. Apetale; 3. Gamopetale (the Monopetale
of Jussieu better named): 4. Dialypetale (the Polypetale of
Jussieu, &e.). The cohort in Endlicher’s classification, it will
be seen, is a higher grade than that to which this name was
applied by Lindley in the more recent use. For the latter, 7. e.
for the grade between these and the order, Endlicher employed
the name of class.

695. Finally, the Genera Plantarum, now in course of pub-
lication by George Bentham and Joseph Dalton Hooker, adopts
in a general way the Candollean sequence of orders, with vari-
ous emendations ; divides the class of Dicotyledons into two sub-
classes, Angiospermous and Gymnospermous; the former into
the Polypetalous, Gamopetalous, and Apetalous divisions ; and
the first of these into the Thalamiflorous, Disciflorous, and Caly-
ciflorous ‘‘ series ” (the middle one composed of the latter part

342 TAXONOMY.

of DeCandolle’s Thalamiflore with some of his Calyciflore) ;
and under these the orders are arranged in cohorts, — fifteen
cohorts in the Polypetale, and ten under three ‘‘ series ” in the
Gamopetale. The remainder of this particular classification
has not yet appeared in print, although partly sketched by its
authors. It will generally be adopted in this country, with some
occasional minor modifications.

696. Various modifications have been from time to time pro-
posed. One of the best of them in principle is that initiated by
Adolphe Brongniart and adopted by many European botanists,
which, recognizing that most apetalous flowers are reductions or
degradations of polypetalous types, intercalates the Apetale or
Monochlamydez among the Polypetalz. But this has never yet
been done in a satisfactory manner, or without sundering orders
which should stand in contiguity.

697. It should be borne in mind that the natural system of
botany is natural only in the constitution of its genera, tribes,
orders, &c., and in its grand divisions; that its cohorts and the
like are as yet only tentative groupings; and that the putting
together of any or all these parts in a system, and especially
in a lineal order, necessary as a lineal arrangement is, must
needs be largely artificial. So that even the best perfected
arrangements must always fail to give of themselves more than
an imperfect and considerably distorted reflection of the plan of
the vegetable kingdom, or even of our knowledge of it.?

1 In the first place, the relationships of any group cannot always be
rightly estimated before all its members are known and their whole struct-
ure understood ; so that the views of botanists are liable to be modified with
the discoveries of every year. The discovery of a single plant, or of a point
of structure before misunderstood, has sometimes changed materially the
position of a considerable group in the system, and minor alterations are
continually made by our increasing knowledge. Then the groups which we
recognize, and distinguish as genera, tribes, orders, &c., are not always, and
perhaps not generally, completely circumscribed in nature, as we are obliged
to assume them to be in our classification. This might be expected from
the nature of the case. For the naturalist’s groups, of whatever grade, are
not realities, but ideas. Their consideration involves questions, not of things,
between which absolute distinctions might be drawn, but of degrees of resem-
blance, which may be expected to present infinite gradations. Besides, al-
though the grades of affinity among species are most various, if not wholly
indefinite, the naturalist reduces them all to a few, and treats his genera,
tribes, &c., as equal units, or as distinguished by characters of about equal
value throughout,— which is far from being the case. And in his works
he is obliged to arrange the groups he recognizes in a lineal series; but
each genus or order, &c., is very often about equally related to three or four
others: so that only a part of the relationship of plants can in any way be
indicated by a lineal arrangement.

BOTANICAL CLASSIFICATION. 343

698. Even the great classes cannot be arranged in a single
line, beginning with the highest Phanogams and leaving the
lowest in contiguity with the higher Cryptogams. The Dicoty-
ledons take precedence of the Monocotyledons in rank. Yeta
part of them, the Gymnosperms, are much the lowest of all
known Phenogams as regards simplicity of floral structure ;
and through them only is a connection with the higher Cryp-
togams to be traced. The Monocotyledons stand upon an iso-
lated side line, and have no such simplified representatives.
In placing the latter class between the Dicotyledons and the
Acrogens, the chain of affinities is widely sundered. If, yield-
ing to a recent 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 Gymno-
sperms to Angiosperms through Gnetaceze and Loranthacez is
not respected. (606, &c.)

699. Nor can the angiospermous Dicotyledons be disposed
lineally according to rank. The apetalous and achlamydecus
must be the lowest. Some are evidently reduced forms of Poly-
petale or even of Gamopetalze: the greater part cannot without
violence be thrust into their ranks. The Gamopetal, especially
those with much floral adnation, should represent the highest
type, the organs being at the same time complete and most dif-
ferentiated from the foliar state. If a natural series could be
formed, these would claim the highest place, with the Composite
perhaps at their head. In the Candollean sequence, they occupy
the middle; and the series begins, not without plausible reason,
with orders having generally complete blossoms, and such as
most freely and obviously manifest the homology of their organs
with leaves, then rises to those of greater and greater combi-
nation and complexity. and ends with those plants which. with
all their known relatives, are most degraded or simplified by
abortions and suppressions of parts which are represented in the
complete flower. These are low in structure, equally whether
we regard them as reduced forms of higher types, or as forms
which have never attained the full development and diversifica-
tion which distinguish the nobler orders.

700. Actual classifications, in their leading features and in
their extension to the cohorts, orders, &c., must be studied in
the systematic works where they are brought into use. In
these are the applications of the principles which are here
outlined. A separate volume of this text-book should illustrate
the structure, relations, and most important products of the
phznogamous natural orders, as another is to illustrate the

344 TAXONOMY.

cryptogamous orders. A synoptical view of the great divi-
sions only, as at present received and named, is appended.
Definitions and characters may be sought in the present and
preceding chapters.

SERIES I. PHAENOGAMOUS OR FLOWERING PLANTS.

Crass L DICOTYLEDONS.

Supcrass I. ANGIOSPERMS.
Div. 1. POLtyPpetTaLous.
Div. 2. GAmMopetTatous. (Monopetalous.)
Div 3. APETALOUS.

Suscrass Il. GYMNOSPERMS.
Crass Il. MONOCOTYLEDONS.
Div. 1. Spapicrovs.

Div. 2. PETALOIDEOUS.
Div. 3. GLUMACEOUS.

SERIES II. CRYPTOGAMOUS OR FLOWERLESS PLANTS.

Crass III. ACROGENS.
Div 1. Vascurar. (Ferns and their allies.)
Div. 2. CELLULAR. (Mosses and Liverworts.)

Crass IV. THALLOGENS OR THALLOPHYTES.

PHYTOGRAPHY. 345

CHAPTER X.
PHYTOGRAPHY.

Section I. NOMENCLATURE.

701. Payrocrapny is the department of botany which relates to
the description of plants. ‘This includes names and terms, also
figures and signs, as well as characters and detailed descriptions.
It comprises two sorts of names, one used to designate organs
or modifications of organs, the other to distinguish plants or
groups of plants. The former is Glossology or (to use the more
common but less proper word) Terminology. The latter is prop-
erly Nomenclature. |

702. Names of Plants were at first only generic names. The
language of botany being Latin, and the plants which the old
herbalists knew being mostly European, their scientific names
were mainly adopted from the ancient Romans or, through Latin
literature, from the Greeks. Ex. Quercus, Prunus, Rosa, Rubus,
Trifolium ; and of Latinized Greek names, Agrostis, Aristolochia,
Colchicum, Melilotus. To the classical names others were
added from time to time ; as, from the Latin, Bidens, Convallaria,
Dentaria ; from the Greek, Anacardium, Glycyrrhiza, Loranthus,
&e. Some barbarous or outlandish names were early adopted,
such as Alhagi from the Arabs, and Adhatoda and Nelumbo from
India. These are mostly such as were or could be conformed to
Latin; as Datura and Ribes from the Arabic, and later 7hea and
Coffea. Of American aboriginal names, Hura, Guaiacum, and
Yucea are examples. Some ancient names of plants commem-
orated distinguished men. Ex. Asclepias, Euphorbia, Lysimachia,
Peonia. Tournefort and his contemporaries resumed this
practice, and named plants in memory or in honor of distin-
guished botanists. Ex. Begonia, Bignonia, Cesalpinia, Fuchsia,
Gerardia, Lobelia; Lonicera, Magnolia.

703. When among plants of the same name or kind different
species were known, these were distinguished by annexed epi-
thets. For example, among the Pines there were: Pinus syl-
vestris, vulgaris ; Pinus sylvestris, montana altera ; Pinus sylvestris,
montana tertia; Pinus sylvestris maritima, conis firmiter ramis
adherentibus ; Pinus maritima minor ; Pinus maritima altera, &c.

346 PHYTOGRAPHY.

And as the number of known species increased, so did the
length of the phrases which were needed for their discrimina-
tion. These ‘* differentia,” thus used as specific names (the
nomina specifica of Linnzeus), became extremely cumbrous. It
was about in the middle of his career that Linnzus suggested
what he called frivial names (nomina trivialia) for the specific
name, consisting of a single word; and in the Species Plan-
tarum, in 1753, he carried this idea into full effect in Botany.
The step was a simple one, but most important; and Linnzeus
himself. who generally did not underrate his services to science,
seems hardly to have appreciated its practical value.*

704. The Binomial Nomenclature in Natural History, thus
established, first separated the name of a plant or of an animal
from its diagnosis, descriptive phrase, or character, and reduced
the appellation to two words, the first that of its genus, the sec-
ond that of its species. The generic name very nearly answers to
the surname of a person, as Grown or Jones; the specific answers
to the baptismal name, as John or James. Thus, Quercus alba is
the botanical appellation of the White Oak; Quwereus being that
of the genus, and ala (white) that of a particular species ; while
the Red Oak is named Quercus rubra ; the Scarlet Oak, Quercus
coccinea; the Live Oak, Quercus virens; the Bur Oak, Quercus
macrocurpa: Magnolia grandiflvra is Large-flowered Magnolia ;
M. macrophylla, Long-leaved Magnolia, and so on. The name of
the genus is a substantive, or at least is a word taken as a substan-
tive. That of a species is mostly an adjective adjunct, always
following the generic name and in the same gender.?. This com-
bination of generic and specific name is the name of the plant.

705. By this system, not only is the name of the plant reduced
to two words, but a comparatively moderate number of words
serves for the complete designation of more than 120,000 plants,*

1 Moreover, he may be said to have adopted rather than originated the
idea; for single-worded specific names were used half a century previous
by Bachmann, a/ias Rivinus.

2 It is to be noted that the classical Latin names of trees are all feminine,
therefore Quercus alba, Pinus rigida, &e.

8 The name of a Sean. is sometimes written in between the two parts
of the plant’s name, as Prunus (Padus) Virginiana. This is the name of the
plant and something more. In addition to the name of the species, that of
the variety or even subvaricty is sometimes added.

# Alphonse DeCandolle several years ago estimated the known species of
Flowering Plants at between 100,000 and 120,000. The larger number may
perhaps include the higher orders of the Flowerless series. In the present
state of our knowledge of the lower orders of Cryptogams, no close estimate
can be well formed of the actual number of species.

NOMENCLATURE. S47

in a manner which avoids confusion and need not overburden the
memory. The generic part of the name is peculiar to each
genus. The specific adjunct is not available for more than one
species of the same genus, but may be used in any other genus.
They are so widely thus employed that the number of specific
may not exceed that of generic appellations.

706. To render this system of nomenclature most serviceable
for the ready identification of such numbers of plants or groups,
and for the clear and succinct presentation of or reference to
what is known and recorded of them, rules are indispensable,
and conformity to admitted rules is a manifest duty. Such rules
were systematically formulated first by Linnzeus, in his Funda-
menta, Critica, and Philosophia Botanica, chiefly for generic
names, some of them being of the nature of laws, some rather of
recommendations. The most important of them remain in full
force, while many of the more particular rules restricting the
choice of names have been abandoned. The code was judi-
ciously revised (in his Théorie Elémentaire) by DeCandolle
‘* who was ruled by the idea of having the law of priority prop-
erly respected,” was critically considered by Lindley in his In-
troduction to Botany, and has of late been reformulated by
Alphonse DeCandolle under the sanction of a Botanical Congress
held at Paris in 1867.!

707. Rules for Naming Plants. These ‘‘should neither be
arbitrary nor imposed 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

1 Lois de la Nomenclature Botanique, ete., Geneva and Paris, 1867. In
the English edition, translated by Weddell: Laws of Botanical Nomenclature
adopted by the International Botanical Congress held at Paris in August,
1867, together with an Historical Introduction and a Commentary, London,
Reeve & Co., 1868. The Laws, simply, were reprinted in the American Journal
of Science and Arts, July, 1868. A few special points have been more recently
discussed by various critics, especially in the Bulletin of the Botanical Society
of France, and in that of the Royal Botanical Society of Belgium. See like-
wise American Journal of Science and Arts for September, 1870, and August,
1877 ; also, Bentham in Journal of the Linnean Society, xvii. 189-198, in which
a just distinction is indicated between changing a well-established name and
giving a new name to a new plant. See American Journal of Science for
April, 1879.

Mention should also be made of Strickland’s Report of a Committee on
Nomenclature to the British Association in 1842, of Agassiz’s classical preface
on the nomenclature of genera in his Nomenclator Zoologicus, and of Dall’s
thorough and well-digested Report of the Committee on Zoological Nomen-
clature to the American Association for the Advancement of Science, 1877,
— these dealing primarily with zoology.

348 PHYTOGRAPHY.

that may create error or ambiguity, or throw confusion into
science. Next in importance is the avoidance of any useless
introduction of new names. Other considerations, such as
absolute grammatical correctness, regularity or euphony of
names, a more or less prevailing custom, respect for persons,
&c., notwithstanding their undeniable importance, are relatively
accessory.” (Alph. DeCandolle, |. ¢.)

708. The following are universal rules in scientific nomen-
clature : —

1. Names must be in Latin or be Latinized. Those from the
Greek (which are more and more abundant, owing to the facility
of this language for compounding) take Latin form and termina-
tion. Those from modern or other than classical languages
should at least have a Latin termination.2 Hybrid names,
namely, those formed by the combination of two languages (at
least of Latin and Greek), should not be made.? ;

2. For each plant or group there can be only one valid name,
and that always the most ancient, if it is tenable.

3. Consequently, no new name should be given to an old plant
or group, except for necessity. That a name may be bettered
is no valid reason for changing it.

709. Names of Genera are substantive and singular, of one
word; and the same name cannot be used for two genera of
plants.* They may be derived from any source whatever, from

1 Thus, words ending with the Greek os generally change it to us, and with
ontoum. <A rule not always observed; for while we have Epidendrum and
Oxydendrum, Linneus himself variously wrote Liriodendrum and Liriodendron,
Rhododendrum and Rhododendron ; and the Greek form now prevails.

2 In this as in other cases, some exceptions are well established by
custom, but they ought not to be extended. The rule as to Latinization is
restricted as respects orthography by the necessity of preserving modern
commemorative names in a recognizable form.

3 But we cannot change numerous old names for this fault, such as con-
voluloides, fumario‘des, ranunculoides, and scirpoides (though they ought to
have been convolrulina, ranunculina, and scirpina) ; and modern botanists have
not scrupled to append the expressive and convenient Greek term -o/des (sig-
nifying likeness) to generic names not of classical origin. Ex. abutiloides,
biroides, davallioides, fuchsioides, gentiunoides, lobelioides, tournefortioid:s. In Eng-
lish, some hybrids will perpetuate themselves, as for instance lerminology,
centimetre, millimetre, beaurocracy, &e.

4 Very many, indeed, are adjectives used as substantives, as Arenaria,
Clararia, Saponaria, Impatiens, Trientalis, and even Gloriosa, Mirabilis, &e.

Some two-worded generic names anterior to Linnzus, such as Dens Leonis,
Vitis Idea, Bursa Pastoris, remain for sections and species, but not for gen-
era. When two words are confluent into one, they are not objectionable,
as Laurocerasus, Carlemania (commemorating Charles Leman, Carolus Le-
manus), &c.

NOMENCLATURE. 3840

prominent or peculiar character or appearance, from localities,
from the names of persons (especially of discoverers), from
indigenous or vulgar names, or even from arbitrary combins-
tions of letters. Unmeaning names, if not in principle the best,
are never misleading. ‘The main requisite is that they should
be euphonious, not too long, and that they should be adaptable
to the Latin tongue. Characteristic names, when possible, are
among the best; such as Sungucnaria for an herb with red juice,
Hemutoxylon for the Logwood tree, Lithospermum for a plant
with stony seeds (or seeming seeds), Myvsurus for a plant with
gyneecium resembling the tail of a mouse. Names of this sort
do not always hold out well; for Chrysanthemum, so called from
its golden yellow blossoms, now has many white-flowered species,
Polygala is wholly destitute of milk, and many species of Con-
volvulus do not twine. Neat anagrams are not bad, such as
Brown’s 7el/ima for a genus nearly related to Mitella. Personal
generic names are wholly proper when dedicated to botanists,
especially to the discoverer of the plant, or to other naturalists,
or to persons who have furthered botanical investigation or
exploration. Ancient names of this kind have been mentioned,
also some of those which commemorate the earlier botanists.
(702.) At present, almost every devotee of the science is thus
commemorated, from Linnzeus and Jussieu downward. In
forming such names, the name of the person, cleared of titles
and accessory particles (thus Cundollea, not D.candollea), takes
the final -a or-/a and becomes feminine ; and its orthography is
preserved as far as possible, making only necessary concessions
to euphony and to the genius of the Latin language.?

The Linnean canon forbade the use of the same generic name in botany
and zoology, — a rule now impossible to maintain. Perhaps we cannot pre-
vent the duplication of phenogamous names in the lower Cryptogamia.

1 Thus, we may write Lescuria instead of Lesquereurra, although Michauria
is the form for the genus dedicated to Michaux, however pronounced. The
genus dedicated to Strangways is written Stranvesia (although Strangwaysia
might have been tolerable) ; to Andrzeiowsky, Andreoskia ; to Leeuwenhoek,
Levenhookia (although the elder DeCandolle restored all the vowels), &e.
As specimens of overdone simplification, there is Gundelia, named for Gun-
delsheimer, and Goodenia, named for Bishop Goodenough, although Gundels-
heimera would not in these days be objected to, and Goodenovia is faultless.
Yet the names having been so introduced into the science should remain,
fixity being of more importance than perfection. Mistaken orthography
of the name itself may, however, be set right. Brown’s Lechenaultia is Les-
chenaultia, Nuttall’s Wisteria (named after Dr. Wistar) is Wistaria. The
rule laid down in the code as drawn up by Alphonse DeCandolle is:
“When a name is drawn from a modern language, it is to be maintained
just as it was made, even in the case of the spelling having been misunder-

350 PHYTOGRAPHY.

710. The etymology of a new genus should always be given.
Of the Linnzan restrictions, one holds, viz. that the names of
genera are not to end in -odds, as many of the older names did.

711. Names of Species are commonly and by preference adjec-
tives, agreeing with the name of the genus, and expressive of
some character, habit, mode or place of growth, time of flower-
ing, or commemorating the discoverer, first describer, or some
one otherwise connected with its history. Thus, in the genus
Ranunculus, R. bulbosus is named from the bulb-like crown or base
of the stem; &. acris, trom the acridity of the juice; J. scede-
ratus (the accursed), in reference to the same property ; 2. repens,
from the creeping habit of the stems; &. pusilius, from general
insignificance ; /. aquatilis, from its growing in water; R. ni-
valis, from living near eternal snow; /. Pennsylvanicus, from
country or State whence it was first made known to botanists ;
fh. Bonplandianus, in honor of Bonpland, one of the discoverers ;
and so on. More commonly, when a discoverer or investigator
of a species is commemorated in the name, this is a substantive,
in the genitive, as Ranunculus Nuttaliii, ¢. e. the Ranunculus of
Nuttall, instead of 2. Nuttallianus, the Nuttallian Ranunculus.
Yet the latter form is preferred when the species is named in
honor of some one who did not discover nor treat of it (which
should seldom be) ; but this distinction is a custom rather than
a rule, and the form of the commemorative name may be settled
by euphony or convenience. In any case, the personal name
should have a capital initial.

712. Many specific names are substantives, occasionally a
common substantive, as Stellar/a nemorum (of the groves), Con-
volvulus sepium (of the hedges), Cassia pumilio (the dwarf ) ;
more commonly it is a substantive proper name, and this usu-
ally an old generic name reduced to that of a species. Ex.
Ranunculus Flammula, R. Thora, and R. Cymbalaria; also Lirio-

stood by the author, and justly deserving to be criticised.” But this is
somewhat too absolute, since it is allowed that obvious errors in the con-
struction of names of Latin or Greek derivation may be corrected, provided
the change does not affect the initial letter or syllable, and that no ancient
names are to be disturbed.

The clause that forbids changes in the orthography of ancient names,
even to make them classical, is a very proper one. The botanical Latin
of Tournefort, Linneus, Jussieu, and their contemporaries, has by pre-
scription rights which botanists are bound to respect. Wherefore Pyrus
is the botanical name of the pear-tree, notwithstanding the classical Pirus.
So Jervis, as a specific name for a smooth plant (and as distinguished from
lévis, a light or slight one), is fixed by long botanical use, although only
lévis is classical; and it is unnecessary to change Ranunculus acris to R. acer.

NOMENCLATURE. 351

dendron Tulipifera, Rhus Toxicodendron, Dictamnus Fraxinella.
These proper specific names take a capital initial letter.1 Rarely
such a name is in the genitive ; as Heterotheca Chrysopsidis, mean-
ing a species of Heterotheca with the aspect of a Chrysopsis.

713. Specific names should be of a single word. Some few
are compounded, as purpureo-caruleum; and some of. ancient
origin (once quasi-generic) are of two words. Ex. Panicum
Crus galli, Cupsella Bursa pastoris, Taraxacum Dens leonis.

714. A specific name cannot stand alone. It is nothing
except as connected with the genus to which it pertains. A
Japonica by itself is wholly meaningless. A plant is named by
the mention of its generic appellation followed 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 FR. agquatilis, var. tricho-
plyllus. Varieties of low grade need not be named. They may
be designated by numbers, or by the small letters of the Greek
alphabet, «, p, &c. When the varieties are marked « and B,
the first is supposed to be the type of the species, or both to
be equally included in the common character. But when the «
is not used, the varieties rank as deviations from the assumed
type of the species. Varieties of cultivation, half-breeds or
cross-breeds, and the like, should have only vernacular names,
at least not Latin ones such as may be confounded with true
botanical names.

716. Names of Hybrids are difficult to settle upon any com-
plete system. When of unknown or uncertain parentage, they
have been named in the manner of species, but distinguished by
the sign & prefixed. Ex. & Salix capreola. Hybrids of known
parentage are named by combining the names of the two pa-
rents, thus: S. purpureo X daphnoides, or K S. purpureo-daph-
noides, for a cross between S. purpurea and 8. daphnoides, of
which the first supplied the pollen to fertilize the second. The
counterpart hybrid is & S. daphnoideo-purpurea.

1 In respect to the initial of geographical specific names, being adjec-
tives, such as Americana, Canadensis, Virginiana, Europea, Anglica, usage
governs, and this is divided. But the elder DeCandolle, who ruled in all
such matters in the preceding generation, always employed the capital in-
itial, and two generations of DeCandolle follow the example. Most English
authors until recently and some continental ones adopt this usage; and it
accords with the genius of the English language, in which we always write
European, British, American, &c., with a capital initial. Of late it is a usua!
practice to write such geographical specific names with a small initial.

b02 PHYTOGRAPHY.

717. The Fixation and Precision of Names. The name of a
plant is fixed by publication, and takes its date from the time
when it is thus made known to botanists.

718. A genus or other group is published when its name
and characters (or the differences between it and all other such
groups) are printed in some book, journal, or other adequate
vehicle of publication, which is placed on public sale, or in
some equivalent way is distributed among or within the reach
of botanists. A printed name without characters, and charac-
ters without name, do not amount to publication.!

719. A species is not named unless it has assigned to it both
a generic and a specific name. It is not published until it is
made known, by name and characters (or by name along with
sufficient information as to its characteristics), in the manner
aforesaid. (718.) Adequate distribution, among botanists and
public herbaria, by sale or otherwise, of a collector’s or distrib-
utor’s specimens, accompanied by printed or autograph tickets,
bearing the date of the sale or distribution (that is, publication
by named /xrsiccate 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 phytography,
not to nomenclature ; but proper identification of names requires
that the name of the author and the time and medium of pub-
lication should be taken into account. Anterior to the binomial
nomenclature, the botanical name of the common tall Buttercup
was ‘* Rununculus pratensis erectus acris,” according to Bauhin,
in his Pinax, p. 179. Under the new nomenclature, which re-
duced the specific part of the plant’s name to one word, this
became Ranunculus acris in Linnzeus, Species Plantarum (ed. 1),
p- 904; and a brief character gave its distinctions. In later
works it has been more fully dcseribed, in some illustrated by
figures. The citation of these works arranged in chronological
order (or in some order), with reference to volume, page, and
in some cases figures, is the bibliography of the plant.? A bot-

' Names may be communicated, in manuscript or otherwise, by the pro-
pounder to an author who may make them known by publication; but the
date of the genus or other group is that of actual publication.

2 This does not cover all the conditions of publication, since it does not
specify the characters (and the same may be said of a published figure,
with analyses) ; but, on the other hand, it conveys to the competent person
receiving the same all this information and more: so that it should carry
the rights of true publication as against any author to whom such names
are or should be known. That is, such are not in the category of “ unpub-
lished names,” which generally ought to be left untouched.

8 For good examples of bibliogranhy, see such detailed works as De-

NOMENCLATURE. 30e

anist, in referring to this or any other plant, might cite any work
which describes it, or none at all. Ranunculus aeris by itself,
as it happens, would lead to no ambiguity. Not so with many
names. For the accurate indication of the species, it is generally
needful, or highly convenient, to specify at least the name of
the author who first published the adopted appellation. So we
write Ranunculus acris, Linn., or L., the abbreviated name of
Linnzeus.! Here we have the name of the plant, and the bibli-
ography reduced to its initial. To this, further citation and
other references may be added or not, as the particular case
requires. But, so far as citation or reference proceeds, it should
simply state the history correctly and clearly.

721. When a species is said to be of Linnzus or DeCandolle
or Bentham, it is simply meant that the adopted name of the
plant (consisting of the generic and specific parts together). was
first published by this author. Some other author may have
named it differently, and even earlier. The earlier name may
have been discarded because the specific portion of it was un-
tenable, either on account of preoccupation or for other valid
reasons. Or the later author may have differed from the earlier
in his views, and have referred the plant to some other genus.
As instances of the first, Huphorbia 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 naine of a different species of the south
of Europe. Whereupon, as the North American species had
no other trivial name, a new one had to be given to it; and it
was named 7. Darlingtonit, in honor of the discoverer and first
describer. The common Milkweed of Atlantic North America
was named by Linnweus Asclepias Syriaca. As this plant is not
indigenous to any part of the Old World, and does not at all
inhabit Syria, this trivial name is not merely faulty but false ;
so it was changed by Decaisne into A. Cornuti, in commemora-
tion of an ante-Linnzean botanist who collected it in Canada and
gave the first account and figure of it. As an instance of the
second, take the pretty little vernal plant Anemone thalictroides,
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 first
done in Michaux’s Flora; and so the accepted name is Thalictrum

Candolle’s Systema Vegetabilium, and Sereno Watson’s Bibliographical
Index to North American Botany, in the Smithsonian Miscel!aneous Col-
lections.

1 For Abbreviations of Authors’ Names, see 385.

304 PHYTOGRAPHY.

anemonotdes, Michx., meaning an Anemone-like Thalictrum, and
Michaux is the authority for this name. The names which for
any reason are superseded become Synonyms? (75d.)

722. A later author may circumscribe a species or a genus
differently from the originator of the name. To a greater or
less extent, this must continually happen in the course of time.
But ** Ricinus communis, Linn.,” stands unmoved by the sub-
sequent admission of various species (known or unknown to
Linneeus) and the final reduction of all to one by a thorough
monographer. So does Silene Gallica, Linn., although S. guin-
guevulnera, Linn., of the same date, is reduced to it. There is
no suflicient reason for writing Myosotis, Brown, or Cynoglossum,
Brown, because this author restricted the limits of these genera ;
nor to write Gilia, LBenth., because Bentham vastly extended

1 The synonymy is an essential part of the bibliography or scientific
history of a genus or species. But synonymous and admitted names ought
to be kept distinct. Keeping this principle in view, —also the decisively
affirmed doctrine of the founder of our nomenclature, that the specific name
is a nullity apart from the generic (so that only the combination of the two
makes the name of the plant, as truly as the constituent halves make the
scissors), and bearing in mind the fundamental importance and absolute-
ness of the rule that no new names ought to be made where there are tena-
ble old ones, —the student need not be misled by the confusing (however
specious) innovation countenanced by many zoologists and some botanists,
and which has of late years been very fully discussed.

The true rule is: “ For the indication of the name or names of any group
to be accurate and complete, it is necessary to quote the author who first
published the name or combination of names in question.” (A. DC.) Thus,
Leontice thalictroides, Linn., fulfils the condition, except where a reference to
the work as well as the name of the originator of the name is demanded.
Then the citation would continue, “ Spec. P/. 312,” and might be further ex-
tended. In the Flora of Michaux, this plant was treated as distinct from
Leontice in genus; and some botanists adopted this view, while others of
equal authority did not. Those who adopt Michaux’s genus name the plant
Caulophyllum thalictroides, Michz.

Now some naturalists quote for the species the author who originated
the trivial appellation even when transferred to another genus. They
would adopt the genus Caulophyllum, yet write: Caulophyllum thalictroides,
Linn. Or else they would avoid direct falsification of the facts by adding
(sp.), this being explained to mean that the specific part of the name only
was given by Linneus. Then, as this omits all mention of the original gen-
eric part of the name, others add this ina parenthesis, and write: “ Caulo-
phyllum thalictroides (Linn. sub Leontice) Michr.,” or “ Caulophyllum (Michx.)
thalictroides, Linn. sub Leontice,” or “Caulophyllum (Leontice, Linn.) thalictroides,
Michr.”’ All such endeavors to mix synonymy with nomenclature appear
to be faulty in principle and unwieldy in practice. In the most abbreviated
form, they state that which is not true: in the others, they impair the sim-
plicity and brevity of the binomial nomenclature. It is all but certain that,
if the genus Caulophyllum had been published in the lifetime of Linnzus,
he would not kave adopted it.

NOMENCLATURE. 355

the comprehension of this genus. Yet in their proper place
such changes may be indicated by ‘‘ pro parte,” or ‘* char. muta-
tis,” ‘‘ excl. sp.,” and the like, — useful qualifying statements,
but no part of the name.

723. Exactness requires that when a group is changed from
a higher to a lower rank, or the opposite, the name of the
author who made the change should be quoted.’ He alone is
responsible for it. But this rule has only recently been strictly
observed.

724. In transferring a species from one genus to another, its
specific name must be preserved (with alteration of the gender,
if need be), unless there is cogent reason to the contrary. It
must necessarily be changed when there is already in that genus
a species of the same name; and then synonymous names of
the transferred species have their claim in order of date. But
whatever name is first 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.?

1 Thus, Potentilla Canadensis, L., var. simpler, Torr. § Gray, and not of
Michaux, for it is the species P. simpler, Michr. Geum, subgen. Stylipus,
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, Uriseb ;” for even if it exactly corresponded with
Cytisus sect. Laburnum of DeCandolle, the latter is not a group of equiva-
lent rank.

But, as to genera and subgenera, this precision should not be insisted on
for times quite anterior to the recognition of such rules and of their need.
Sperguaria began with Persoon as a subgenus in the year 1805, and this
date has been assigned to the genus, although it was taken up as such only
in 1819 by Presl and in 1824 by Bartling.

2 Thus, in the case of an older specific name being known, as that of
Chilopsis saligna, Don, recognized as Bignonia linearis, Cav., though Don ought
to have adopted the latter trivial name, yet as he did not (and the rule was
not then really in force as now), there was no need for the introduction
of a third name, Chilopsis linearis, DC. “So, again, an Indian Grass was
first named and described by Willdenow as Coir arundinacea, then named
by Roxburgh as Coir larbata, and entered in Sprengel’s Systema with
Willdenow’s character as Coir Kenigii. All these names were defective
as referring to a wrong genus. Brown corrected the error by creating the
new genus Chionachne, and selected Roxburgh’s specific name as the one
most generally known and the least liable to misinterpretation , and Brown’s
Chionachne barbata is therefore the first correct name; for which Thwaites
afterwards substituted Chionachne Kanigii, 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-
acez, in Jour. Linn. Society, xvii. 197, 198, November, 1878). The following

3856 PHYTOGRAPHY.

725. Names of Subgenera or of other sections of genera are
like those of genera; indeed very many of them, and the most
fitting, are eld generic names which haye been comprehended
in the genus by reduction. Unlike genera and higher groups,
however, sections, when of Greek derivation, may properly take
the termination in -ovdes,t and the typical section may bear the
name of the genus with the prefix Lu.” Sections need not be
named at all, and only those of comparatively high rank should

is a farther extract from the same protest against the practice ‘of creating
a new name in order to combine an old specific with a new generic one: ”
“In Ferns, the wanton multiplication of ill-defined or undefinable genera,
according to the varied fancies of special botanists, has had the effect of
placing the same species successively in several, sometimes seven or eight,
different genera ; and it is proposed to maintain for the specific appellation
the right of priority, not only in the genus alone in which it is placed, but
in the whole of the genera to which, rightly or wrongly, it has been referred.
This has been carried to such an extent as to give to the specific name a
general substantive aspect, as if the generic ones were mere adjuncts, —
a serious encroachment on the beautiful simplicity of the Linnean nomen-
clature; and it is to be feared that there is a tendency in that direction in
phenogamic botany. When a botanist dismembers an old genus, the rule
requires that he should strictly preserve the old specific names in his new
genera; and, when he has wantonly and knowingly neglected this rule, it
may be right to correct him. But where a botanist has established what
he believes to be a new species, and has therefore given it a new name, the
changing of this name after it has got into general circulation, because it
has been discovered that some other botanist had previously published it
in a wrong genus, is only adding a synonym without any advantage what-
ever, and is not even restoring an old name; for the specific adjective is
not of itself the name of a plant. ... A generic name is sufficiently in-
dicated by one substantive; for no two genera in the vegetable kingdom
are allowed to have the same name; but for a species the combination of
substantive and adjective is absolutely necessary, the two-worded specific
name is one and indivisible ; and combining the substantive of one with the
adjective of another is not preserving either of them, but creates an abso-
lutely new name, which ought not to stand unless the previous ones were
vicious in themselves, or preoccupied, or referred to a wrong genus. It is
probably from not perceiving the difference between making and changing
a name that the practice objected to has been adopted by some of the first
among recent botanists.” Bentham, I. e.

1 A genus could not properly have one of its sections called by its own
name with the addition of -o/des or -opsis, as Asteroides or Asteropsis, for it
is senseless to declare that an Aster resembles an Aster; but sectional names
of this composition may be excellent for sections of other genera, as ex-
pressing analogy or resemblance. Latin generic names used for sectional
ones properly take the addition of -e//a, or -ina, or -astrum.

2 The prefix Lu (Greek for much, very, or true), prefixed to a generic
name of Greek origin, is the proper designation of the typical section of
that genus, meaning the group which should bear the generic name if such
genus were divided. The rule against hybrid names should in strictness
exclude this prefix from Latin names, but it has not always done so.

NOMENCLATURE. 307
have substantive names. Designations, however, are conven-
ient for lower sections; and the name of a leading species
may be used, in the plural; as Aster, section Amelli, and sect.
Concinni. Subgenera need not agree in gender with the genus
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 (Malus) coronaria.

726. Names of Tribes, Orders, &c. The names of all groups
superior to genera are adjectives plural, and with few excep-
tions are the names of genera lengthened by some adjective
termination. Ex. From Rosa, Rosee, Rosacee, Rosales; from
Myrtus, Wyrtee, Myrtaceae, Myrtales ; from Berberis, Lerberidee ;
from Tamarix, Zumaricinee ; from Salix, Sulicee, Salicinee.
The substantive Plante being understood, the groups are Rose-
ous, or Rosaceous, or Rosal plants, &e.

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@. Ex. Rosee, Phasevlee, Antirrhinee, Oxalidee,
&¢. Some subtribes take the name of the tribe with the prefix
Eu, as Eiphaseolee tor that subtribe of the tribe Fhaseolez
which comprises the representative genus Phaseolus. Tribal
names may take the same prefix, as Huc@salpinee for the tribe
of the suborder Ceesalpinez which contains the typical genus
Cvesalpinia.

728. Ordinal Names are formed in the same way, but with a
preference for certain terminations which may denote their rank,
especially that of -acee,— as Rosacea, Myrtacee, Cucurbiiacee,
meaning Rosaceous, Myrtaceous, and Cucurbitaceous plants.

729. The names of what we now call natural orders, as
sketched or adopted by Linnaeus, were mostly descriptive, such
as Ensate, Spathacee, Coronarie, Pupilionacee, Conifere, Amen-
tacee, Umbellate ; but a few took their names from genera, as
Orchidee, Liliacee. 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 Leguminose, Corymbifere, &c. But the greater part are
simply plurals of generic names, such as Asparag/, Junci, Lilia,
Muse, Orchides, Lauri, Convoleuli, Erice, Acera, Cacti. To a
few was given the lengthened termination in -ee@, as Polygonee,
Solanee, Berberidee, Curyophyillee ; to some, the termination in
—ace@, as in Cichoracee, Campanulacee, Leubiacee, Ranuncu-
lacee, Malvaceae, Tiliacee, Cucurbitacee. Subsequent authors
have necessarily changed all names which were plurals of gen-

358 PHYTOGRA PHY.

era; and the strongly prevalent tendency has been to give the
termination in -acee to all such ordinal names, and to restrict
this termination to orders. Lindley insisted upon making this .
an absolute rule even for names not formed from generic appel-
lations: but this will not be adopted.

730. In the first place, several large orders which have been
known from the first by such characteristic names as Crucifere,
Leguminose (and its suborder Papilionacee), Guttifere, Umbelli-
fere, Composite, Labiate, Cupulifere, and Conifere, also Palmee
and Graminee, Filices, and even Aroidee and Ficoidee, will retain
these appellations ; but no new ones of the kind will be made.

731. Also, names formed from genera which do not well take
the termination in -ace@ may be allowed as orders to retain their
natural form in -cnee, -idee, -ariew, and the like. Ex. Tamaris-
cinee, Salicinee, Scrophularinee, Berberidee, Lentibuluriee. We
may prefer for the sake of uniformity to write Salicacee, Berberi-
dacee, Lentibulariacee, and Scrophulariacee (as we should write
Violacee), but this form cannot be insisted on. On the other
hand, a termination in -acee has been allowed in the names of
certain tribes to avoid excessive iteration of vowels. Thus, for
the tribe of which Vernonia is the leading genus, authors write
Vernoniacee, to avoid Vernoniee, which ends with four vowels.
Spireea and Staphylea are the types of tribes, for which the names,
if they followed the rule, would be Sprreee and Staphyleee,
ending one in five the other in four consecutive vowels. Some
avoid this by writing Staphyleacee and Spireacee. Others write
Stuphylee, but this is only the plural of the generic name.

752. A few orders or other groups took their names long
ago from superseded generic names. Ex. Caryophyllacee or
Caryophyllee, Onagracee or Onagrariee, and Lentibulariee.

733. Names of Cohorts are distinguished by the termination
in -ales. This was proposed by Lindley, and is adopted by
Bentham and Hooker in the Genera Plantarum. Ex. Ranales,
Parietales, Malvales, Rosales, Passiflorales, &c., most of them
founded on the names of representative genera and orders.
Euphony requires some to take other terminations. Ex. Poly-
galine, Caryophylline.

734. Names of Classes and other great divisions are plurals,
either adjective or adjective nouns, expressive of the leading
character. Ex. Polypetale, Gamopetale, Apetale ; Angiosperme
and Gymnosperme ; Dicotyledones and Monocotyledones. The
names of the two great series or sub-kingdoms, following the
analogy of the Linnzean classes, end in -ia, and are Phenogamia,
or Phanerogamia, and Cryptogamia.

GLOSSOLOGY. 359

Section II. Gtrossotocy or TEermMiInoioey.?

735. This is nomenclature as applied to organs or parts and
their modifications. The actual botanical terminology owes its
excellence in the first place to Linnzeus, and then to DeCandolle.
The Théorie Elémentaire of A. P. DeCandolle (the first edition
of which was published in 1813) is still classical authority, and
until recently has received few additions as regards terms need
ful in phenogamous botany.

736. The fundamental rule is that each organ or part shall
have a substantive name, and that modifications of organs shall
be designated by adjective terms. ‘These names or terms should
be as precise as possible: each object ought to be known by only
one name, yet synonyms are unavoidable; and no term ought
to be used with two different meanings. The word flower, for
instance, must not be used for a cluster of flowers, however it
may imitate the appearance of one, nor for the corolla or other
portions of a flower. Still, some terms have to be used in two or
more senses, to be determined only by the connection, or else as
having both a special and a more general meaning. Leaf ( fo-
lium) is a notable instance. <A bract, to go no farther, is a sort
of leaf; and the imperfect stamens of a Catalpa-flower and
Pentstemon are stamens, although likewise called staminodia:
these are liable to be called sometimes by one, sometimes by the
other name. But, howev er frequent such ambiguities may be in
morphological treatment, they are usually avoidable in descriptive
botany, in which terms are held to their more special or partic-
ular sense. Yet no rule can absolutely determine whether leaf
or bract, bract or bractlet, is the proper term in many cases.
Moreover, substantive names must also be applied to certain
mere modifications of the same organ. In the same family, a
simple carpel, differently modified in fruiting, is an akene ina
Ranunculus, a follicle in Aquilegia, a berry in Hydrastis and
Actzea; while in another family an additional line of dehiscence
makes it a legume. Moreover, in this latter family it is called a
legume when it is not dehiscent at all. and even when it becomes
a drupe! Arbitrary rules cannot absolutely fix technical any
more than ordinary language.

737. Experience and judgment must determine what modifi-
cations of organs should be regarded as a kind, and bear sub-

1 Although the former is the better name, the latter is well established in
use as an English word, and perhaps it need not be objected to, inasmuch as
the Latin terminus comes from the Greek répua, of the same meaning.

260 PHYTOGRAPHY.

stantive instead of merely adjective names. But the former
should not be unnecessarily multiplied.

738. The classical language of scientific botany being Latin,
all the organs of plants and their principal diversities are desig-
nated by a Latin or Latinized name. Modern languages have
also their own names and terms. Greatly to its advantage,
English botanical terminology has adopted and incorporated
terms from the Latin and Greek, with slight changes, not obscur-
ing the identity, thus securing all their precision, and rendering
the simple botanical Latin of descriptions of easy acquisition to
the English student.

739. In a text-book like this, the principal names and terms
applied to organs and their leading modifications, as also those
which relate to their action (physiological terms), or to our study
of them (didactic terms, such as phytography, phyllotaxy, glos-
sology), are defined and illustrated in course. There remain
the more numerous and varied characteristic terms, chiefly adjec-
tives, applicable to more than one or to all organs, and which
compose the greater part of glossology. These, which DeCandolle
arranged systematically with much elaboration, may best be
reached by a glossary or dictionary, such as that at the end of this
volume, which comprises the substantive terms likewise.

740. From characteristic adjective terms are derived the
greater number of specific names of plants ; of which, therefore,
the glossary may elucidate the meaning.

741. Capable as the existing system is, it cannot in single
words define all observed forms and grades, nor well avoid
various ambiguities of meaning. Some defects of the first kind
are remedied by combining with a hyphen two congruous terms
to denote an intermediate state. Ex. ovato-lanceolutus, or ovate-
lanceolate, for an outline between the two. Also a term may be
qualified by the prefix sb, in the sense of somewhat, as in sud-
rotundus, subcordatus (somewhat round or slightly heart-shaped),
or diminutives (such as ¢ntegriusculus), or superlatives (cnteger-
rimus) or Other strengthened forms (such as perangustus) may
be employed. Among terms of more than one form of meaning
are such as calycinus. which may mean, according to the context,
pertaining to the calyx, or of the appearance of calyx; cymosus
may mean in cymes, or bearing cymes, or in the manner of a
cyme; and paleaceus may mean provided or beset with chaff, or
resembling chaff in texture. Often the form of the word should
distinguish the sense ; as fol/atus, furnished with leaves, foliosus,
with abundance of leaves, while foliaceus may mean either bear-
ing leaves, or properly of leaf-like texture or appearance.

DESCRIPTION. 361

742. Absence of an organ or quality may be expressed by
means of a prefix with privative signification, as trdehiscent, not
dehiscent, exannulute, destitute of a ring, apetalous, without
petals. But the Greek privative « should not be prefixed to
Latin words, nor the Latin sud to terms taken from the Greek.

743. When the Latin preposition 04 is prefixed to an adjective
term, it means obyersely ; thus obcordatus is cordate inversed,
that is, the broader end with its notch at the apex (instead of
the base) of the leaf or other plane organ.

Section III. DeEscriIprion.

744. Under this head may be conveniently comprised all that
relates to the form of the exposition, in botanical terms, of the
differences by which the species and groups of plants are distin-
guished and recorded, the structure exemplified, and the history
or bibliography indicated in systematic works or writings. Lin-
nus, in the Philosophia Botanica, treated these topics under
the head of ‘* Adumbrationes.”

745. Descriptions may be full and general, comprising an
account of all that is known of the structure and conformation
of a plant or group, or rather all that is deemed worth recording,
or they may be restricted to what is thought most important. In
the former, the description is independent of all relative knowl-
edge, or takes no notice of relationship to other plants or groups.
The latter intends to portray the species or group in its relations
to others, and to indicate the 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 termed
Characters, or the Character, of the group so described. There
are all gradations in practice between characters and descrip-
tions ; but the distinction should be maintained.

746. Characters are specific, generic, ordinal, &c. They are
the differenti@, ov marks which distinguish a group from any
related group of the same rank with which it may properly be
compared. According to the occasion and purpose, they may
specify 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, orders, &c.*

1 The former would answer to what have been termed d/fferential char-
acters, the latter to essential characters. Linnzeus divided (generic) characters
into factitious, essential, and natural; by the former denoting any difference
which may effectively distinguish between any two groups brought arti-

362 PHYTOGRAPHY-

What is now termed the specific character was the specific name
with Linneus and his predecessors; what we call the specific,
Linneeus called the trivéal name. (703.)

747. Subordination of characters and the avoidance of vain
repetitions require that as far as possible — regard being had to
the form of the work —the ordinal character should contain
only what is needful to circumscribe it, and to exhibit clearly
its morphology ; that the characters of tribes or other divisions
should not reassert any portion of the ordinal character, nor
the generic character that of the superior groups ; and so of the
sections and subdivisions of all grades down to the species.
Equally from the specific character should be excluded every
thing which belongs to the generic, or is common to its rela-
tives generally, or has been already specified in the section or
its subdivisions. So, likewise, of the varicties under the spe-
cies. This can be done only by so arranging the species as best
to exhibit their relationships, that is, by bringing together or
into proximity those of greatest resemblance in all respects,
or in the more important respects. What these are, and how a
just subordination of characters is to be apprehended, cannot
be taught by rules, but must be learned by experience and
from the critical study of the classical botanical works. No one
is competent to describe new plants without such study, and
without a clear conception of the position which a supposed
new species should occupy in its genus, or a genus in its order.

748. Characters of orders, genera, and of all intermediate
groups, are drawn almost without exception from the organs of
fructification. In the description, these parts are mostly taken
‘in order, beginning with the calyx and ending with the ovary,
the fruit, seed, embryo. But, as to the orders, some writers pre-
fer to preface these proper characters with a general sketch of
those derived from the vegetation, which, albeit of less syste-
matic value generally, are often very characteristic of particular
families. Rubiacez, for example, are known by their opposite
entire and simple leaves and intervening stipules, along with a
few floral characters; Sarraceniaceze, by tubular or pitcher-like
leaves, along with a certain combination of a few other charac-

ficially together, as they might be in an artificial key, and as very unlike
genera often were in his sexual system; by the second meaning the distine-
tions, the fewer the better, which will separate a group from its nearest
relatives ; by the third, all real marks of difference, 7. e. all afforded by the
organs of fructification, which only were taken into account for genera, &c.
Upon the construction of this natural character Linneus prided himself,
and justly. These are the characters in his Genera Plantarum.

CHARACTERS. 503

ters, and so on. Where brevity is aimed at, such external and
obvious characters, followed by a few diagnostic marks, may
practically take the place of a full enumeration of particulars,
many of which may be common to other orders, though not in
the same combination. Generic characters always commence
with the calyx or most external of the floral organs and proceed
to the ovary, thence to the fruit and seed, and end with subsi-
diary (but often no less diagnostic) particulars furnished by the
vegetation and mode of growth.

749. Detailed descriptions of species, as distinguished from
technical characters, commence with the root, and proceed in
order to the stem, leaves and their parts or appendages, inflor-
escence, bracts, flowers, calyx, corolla, stamens, with filament,
anther, and pollen, the disk, if any, gyncecium and its parts,
ovules; then the fruit, seed, albumen, if any, embryo and its
parts. But descriptions of this sort in most works and in ordi-
nary cases are partial and subsidiary, comprising only certain
details supplementary to or-in amplification of the character of
the species or genus. In condensed works, such description is
wholly omitted, or is reduced to a few specifications which do not
readily find their way into the character.

750. Specific characters usually follow the same order of
enumeration, from root to seed, so far as the several organs are
mentioned; and in Latin the phrases are expressed in the abla-
tive case. But these particulars are often very conveniently
prefaced by statements applying to the whole plant rather than
to any one organ; and these are given in the nominative, and
agree with the name in gender.

751. Linnzus required that neither the essential character of
a genus, nor a specific character (his nomen specifieum), should
exceed twelve words. Latin characters take fewer words than
English. But this arbitrary rule is wholly out of date. Yet
such characters should be brief and diagnostic: otherwise, their
advantage is lost, and the distinction between them and descrip-
tions disappears. In monographs and floras, the desirable
brevity, or such as the case admits, is secured by proper group-
ing under a subordination of sections, subsections, and other
subdivisions.”

1 Ex. “Nepera Cataria: erecta, elata, cano-pubescens ; foliis petio-
Jatis,’” ete. In English, these adjectives without any substantive expressed
will be seen to belong, as here, to “ plant” or “ herb ” understood.

2 In the Synoptical Flora of North America, such a system of successive
divisions is thoroughly carried out. And, if the specific characters are by no
means short, it is mostly because nearly all separate descriptive matter is

504 PHYTOGRAPHY.

752. Punctuation. In proper descriptions, and in characters
of genera and of higher groups, the account of each organ forms
a separate sentence ; and in Latin the terms are in the nomina-
tive case, except subsidiary portions, which are often thrown
into the ablative. Excepting the latter part, the adjective terms
are separated by commas. <A specific character is always in one
sentence. In Latin, its clauses are mainly in the ablative; and
much diversity prevails as to the punctuation. Subgeneric and
other sectional characters are commonly framed like those of

dispensed with: consequently various particulars are added to the char-
acter which do not strictly belong to it. In Bentham’s great Flora Austra-
liensis, also in English, specific characiers are replaced by a characteristic
synopsis at the head of each genus; and a terse description under each
species completes the account. Moreover, Bentham, in recent works, such
as his revision of the Genus Cassia, also that of the Mimosez, which have
Latin characters, writes these in the nominative case and cach member in a
separate sentence, in the descriptive form, abandoning the long-used abla-
tive form.

1 Linneus employed only the comma in the specific character, along with
a subsidiary use of the colon in a manner very unlike its ordinary use in
punctuation, making it a point of less value than the connna. Thus,

“CHEeNOPoDIUM ALBUM foliis rhomboideo-triangularibus erosis postice
integris : summis oblongis, racemis erectis.” Spec. Pl. ed. 2, 519.

Here, while the two main members of the sentence are separated by a
comma, a subsidiary portion of the first member, relating to the uppermost
leaves, is separated by a colon. Linnzus employed the colon in the same
way in generic characters. This anomalous usage is now abandoned. But
most authors have followed the Linnean pattern in distinguishing the prin-
cipal members by commas only, so that these become the only points in the
specific character, however complicated that may be. Thus,

““Ranuncutus acris (Linn. Spec. 779) foliis pubescentibus subglabrisve
palmato-partitis, lobis inciso-dentatis acutis, summis linearibus, caule erecto
plurifloro subpubescente, pedunculis teretibus, calyce subvilloso, carpellis
mucrone suberecto terminatis ” DeCandolle, Prodromus, i. 36.

This is the punctuation throughout the Prodromus and in most contem-
porary systematic works. Its imperfection is shown in the above-cited speci-
men. The primary members of the sentence, which characterize the leaves,
stem, peduncles, calyx, and carpels, are distinguished by the same grade of
punctuation which serves for the parts of the first member, viz. the lobes
of the leaves, and for a still subordinate portion, viz. the form of the upper-
most lobes. This want of subordination is to be remedied by the use of
semicolons between the principal members, and of the commas only fer the
secondary ones,— a punctuation now not uncommon, and which is adopted
in the recent first volume of the Monographie Phanerogamarum of the De-
Candolles, which supplements the Prodromus. The portion of that volume
contributed by Dr. Masters better exemplifies this than does the rest of the
volume. For the latter sacrifices the advantage of the change by the inser-
tion of commas between each adjective of a continuous abiative phrase (as,
“Smilax laurifolia ; limbis foliorum oblongis vel ovato-oblongis, coriaceis,
3-5-nerviis, subtus pallidioribus,” etc.), where they are generally deemed

PUNCTUATION, SYNONYMY. 369

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 intervene between the specific name and
that of the author cited? The practice varies. But, if the name
is Latin, the comma is superfluous ; for the abbreviated name of
the author is supposed to be in the genitive, and to read thus:
Ranunculus repens Linnei. Still, since when the suthor’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
proper to insert it.

754. In citations, the classical practice is to separate the refer-
ences from each other and from the name by periods; thus,
*¢ Anemone cylindrica, Gray, Ann. N. Y. Lye. 3. 221. Torr. &
Gray, Fl. 1. 113,”? &e. It is becoming equally customary to
separate the several citations by semicolons, thus bringing all the
references under one name into one sentence. The bibliography
of a species or group of species which a describer or other author
has to refer to (with more or less fulness, according to the form
of his work) is to be sought partly under the admitted name,
and partly in the

Zoo. Synonymy. This includes all other than the admitted
names. Ex. Hydropeltis of Michaux is a synonym of Brasenia of
Schreber, the latter being the earlier published name. Nectr/s
of Schreber is a synonym of Cabomba of Aublet, the latter hay-
ing priority. Thalictrum anemonoides of Michaux has for syno-
nyms Anemone thalictroides of Linneus and of many subsequent
authors who followed him in referring this ambiguous plant to
Anemone (721) ; and also Syndesmon thalictroides of Hottmannsegg

5)

and Anemonella thalictroides of Spach, who proposed to consider

superfluous. The preferable punctuation of the character above-quoted
from the Prodromus would be

Ranuncucus acris (Linn.): foliis pubescentibus subglabrisve palmato-
partitis, lobis inciso-dentatis acutis, summis linearibus; caule erecto pluri-
fioro subpubescente; pedunculis teretibus; calyce subvilloso; carpellis
mucrone suberecto terminatis.

The advantages of this style of punctuation will more and more appear,
when applied to less simple cases. Commas between the ablative adjectives
are superfluous and confusing.

In English characters, commas are required between the adjectives which
follow the noun. Rightly to express the subordination of characters, the plan
adopted in the Synoptical Flora of North America is recommended ; that is,
with colons separating the principal members, semicolons for subordinate
and dependent ones, and commas between the adjectives of the same noun.

1 See Watson’s Bibliographical Index to North American Botany (where
this style is adopted) for a general model for the arrangement of synonymy
and citations.

306 PHYTOGRAPHY.

it an intermediate genus between Thalictrum and Anemone. In
systematic works, the specific character immediately follows the
name, and generally forms a part of the same sentence ; and is
followed first by citations of authors who have adopted the name,
and then by the synonymy, or as much of it as the plan of the
work calls for. The synonymous names and the references
under them should be cited in the order of their publication.
But, to economize space, all the authorities for the same name
are brought together into one sentence, and arranged according
to their date. Also, where the synonymy is not elaborately
displayed, the various synonyms of the same generic name are
usually placed in consecutive order.

756. Ieonography. ‘The leading and most essential citation is
that of the author by whom and the work in which a plant is
named and described, and also the work in which it is best char-
acterized." Among the characterizations, published figures hold
a prominent place. The citation of these is an important part
of the synonymy. The best botanical plates are those which
give detailed analyses of the parts of the flower, fruit, and seed,
displaying their structure.

757. Habitat and Station are recorded in a sentence or para-
eraph following the name, character, and synonymy of a species.
The habitation is the place, district, or region at or within which
the plant is known to be indigenous, or to grow spontaneously.
The complete habitat is the geographical range. The station is
the situation it affects, whether in water, in marshes, on shores,
on hills or mountains, in forests, on open plains, &e.

758. Discoverer, &e. To the habitat and station of newly
discovered, rare, or local plants should be appended the name
of the discoverer or the collectors by whom the species has be-
come known to science, at least when the plant is first published.
Date of discovery should also then be indicated.

759. Time of Blossoming should be recorded. either the month
or the season, to which may be added that of the maturity of the
fruit. When the month or season is mentioned without farther
explanation, flowering-time is intended. In a flora, this may
sometimes be indicated under the genus for all the species.
In the flora of an extensive region, and in respect to species of
considerable range in latitude or longitude, the time of flowering
differs so widely at the extremes of the geographical range that
it cannot well be specified except in general terms, as spring,
summer, autumn, &e.

760. Etymology of Names. When a new generic name is pub-
lished, its origin and meaning should always be given, if the

ACCENTUATION, ABBREVIATIONS. 367

nature of the publication will allow it. So likewise-of species,
except where the source or signification of the name is mani-
fest. This is commonly the case as respects most characteristic
specific names, and also those drawn from station, habitat, and
the like.

761. Aecentuation of Names. The pronunciation of botanical
names is settled by the rules of Latin prosody. All that is
usually attempted in those botanical works which take this into
account is to mark the syllable upon which the principal accent
falls. This in, words of two syllables is always the first; in
words of three or more syllables, either the penult (the last sylla-
ble but one) or the antepenult (next preceding syllable). When
the penult is a long syllable, it takes the accent ; when short, this
recedes to the antepenult. The accentuation may accordingly
be sufficiently indicated by marking the quantity of the penult,
either long as in Eyica, or short as in Arbaus and Gludidlus. Or
else the accent may be marked by a proper sign, as Erica, Arbutus,
Gladéolus. An endeavor has been made to represent the longer
sound of the vowel by the grave accent-mark, as Hrica, and the
short by the acute, as Gladiolus. But this plan is encumbered
with practical difficulties. :

762. Abbreviations are required, both of the name of the au-
thor, when of more than one or two syllables, and of the titles
of the works cited. There are also the customary abbreviations
in the citation of volume, page, plate, &c., in which there is
nothing peculiar to botany.

763. The simple rule for the abbreviation of an author’s name
is to abridge it of all but the first syllable and the first letter of the
following one (as Lam. for Lamarck, Hook. for Mooker), or the
first two letters following the vowel when both are consonants
(as Linn. for Linnzeus, Juss. for Jussieu, ich. for Richard).
Sometimes more of the name must be given, in order to distin-
guish those beginning with the same syllable. So we write Michr.
to prevent confusion of the name Michaux with that of Micheli,
which, being the earlier, claims the abbreviation M/ch., and
Bertol. to distinguish Bertoloni from Bertero. Sometimes a-
much-used name of one syllable is abbreviated, as Br. (or R.
Br.) for Robert Brown. Initials or abbreviations of the bap-
tismal name are needed to distinguish botanists of the same
name; as P. Browne in distinction from Robert Brown, Ach.
Rich., Adr. Juss., Alph. DC., to distinguish the younger from the
older Richard, Jussieu, and DeCandolle. Or. where father and
son, the abbreviation for the latter may be Juss. fil., Hook. fil.
or Hook. f., &¢. Certain, but very few, well-known and eminent

268 PHYTOGRAPHY.

names are abbreviated to a sign; as L. for Linnzeus, DC. for
DeCandolle, HBik. for Humboldt, Bonpland, and Kunth, the
latter too long after ordinary abbreviation.’ Care should be
taken to afiix the period by which abbreviations may be dis-
tinguished from full names, such as Don, Ker, blytt.

764. Abbreviations of titles of works follow the same rules as
those of names, or at least are in no wise peculiar in botany.

765. Abbreviations of the names of organs follow the same
rule: Cal. for calyx, Cor. for corolla, Stam. for stamen or
stamina, st. for pistillum or pistil, “r. for fructus or fruit,
Per. for pericarpium or pericarp, Sem. for semen or seed, are the
most common. ab. for habitat or geographical station, Herd.
for herbarium, Gen. for genus, Sp. or Spec. for species, Var. for
variety, and the like, every one will understand. But some
abbreviations which are common in botanical writings, at least
those in Latin, may need explanation to the elementary student.
A list of abbreviations is appended. See p. 390.

766. Signs. Under this head might be ranked such abbrevia-
tions as v. v. for vidi vivam, v.s. for vidi siccam, to note that the
writer has seen the plant, either alive or in a dried specimen ; or,
more particularly, v. s. s., when it is a spontaneous specimen
that has been examined in a dried state, and v. s. e., when it was
a cultivated specimen; v. v. c., when the living plant was seen
in a garden only, and ». v. s., when the spontaneously growing
plant was seen alive. There are also proper signs, of which the
most common are those which indicate the sexes of blossoms,
the duration of a plant, and the like. Also the interrogation
point (?) used to express doubt; the exclamation point (!) to
indicate the certainty that is given by the actual sight of an
authentic original specimen. See p. 391.

767. The marks used to indicate the subordination of sections
under a genus, or in the synoptical arrangement of genera, and
the like, are not settled by any fixed rule. An approved ar-
rangement is to employ the following marks in the given order,
S$ * + + =. The first one, for sections of the highest order,
takes numerals after the sign. Ex. § 1, and so on. When

' As Alph. DeCandolle remarks, the proper abbreviation of the name he
bears is Cand. But the form DC. was very early adopted by the first of
the illustrious name, and has been continued for almost three quarters of a
century. Alphonse DeCandolle would prefer to write it D.C., but has not
adopted that mode, nor should we; for DC. and HBK. are convenient ab-
breviations reduced to signs. But such forms should not be increased. For
ordinary names they would be unintelligible.

Names which are not too long, and of which an abbreviation by the ordi-
nary rule is insufiicient, such as Decaisne, should rather be written in full.

SIGNS, ETC. 369

such sections are followed by a substantive name, they are
equivalent to subgenera.’ Ex. Phacelia, Juss., § 1. Euphacelia,
?.e. the true or typical Phacelia; § 2. Cosmanthus, Gray, Ke.
Sections next in rank to these are marked with asterisks, * for
the first, * * forthesecond, * * * forthe third one of the same
rank. Divisions of these have the + prefixed ; and so on in the
sume way. Still farther subdivisions may be marked by the
small letters of the alphabet consecutively. a, d,c. When capital
letters are used for division marks, it is mostly for those of a
high grade.

768. Floras, Monographs, &e. A systematic work describing
in proper order the plants of a country or district is generally
alled a Flora, A Flora of a small district takes the diminutive
name of a Mlorula. <A universal work of the kind when it ex-
tends to the species is a System, Systema Vegetabilium or Systema
Regni Vegetabilis. The latest completed Systema Vegetabilium
is that of Sprengel (1825-1828), in five octavo volumes, on
a very condensed pian. A compendious Flora or Systema is
often termed a Prodromus, literally meaning a forerunner or
preliminary work. But, as even this is more than most bot-
anists are able to complete, the name of Prodromus is now
applied to works which are not intended to precede fuller ones
by the same author. The principal work of this kind is the Pro-
dromus Syst. Nat. Regni Vegetabilis, commenced by DeCandolle
in the year 1824, continued by his son Alphonse DeCandolle
(aided by various botanists) to its close in 1873, down to vol.
XVii., or essentially twenty very compact octavo volumes, these
carrying the work only through the great class of Dicotyledones.
But the publication of the monocotyledonous orders has com-
menced in a series of Monographs (Monographie Phanerogam-
arum). A Monograph is a systematic account of all the species
of a genus, order, or other detached group.

769. Specimens of botanical characters and descriptions, cita-
tions, &c., illustrating this chapter, might be given here. But,
for those in Latin, the classical works of DeCandolle and others,
and for the genera those of Jussieu, Endlicher, Bentham and
Hooker, may be taken as models. In English, those of the
latter authors, and in the United States the better-known
writings of the present author, especially the later ones, may
be referred to.

1 DeCandolle in the Prodromus employed the word Sect. (Sectio) for
what answers to subgenus or at least to the highest grade of sections ; then
§ 1, § 2, &e., for the next grade below subgenus ; and then the asterisk, and
other marks.

24

370 PHYTOGRAPHY.

Section IV. Specimens; DIRECTIONS FOR THEIR EXAMINA-
TION, PRESERVATION, &¢.

770. Implements. Those necessary for the examination of
phenogamous plants, Ferns, and the like, are a simple pocket
lens, a simple dissecting microscope; also a sharp thin-bladed
knife and some needles of various fineness, mounted in han-
dles, for dissection.

771. Forasingle hand lens, one magnifying only from four to
six diameters is the most useful. A doublet, or a parabolic lens
of Tolles, of about an inch focus, is better, but much more expen-
sive. The simple stage-microscope for dissection need have only
two lenses (doublets or otherwise) with large field and good
definition, one of an inch and the other of about half inch focal
distance ; and a glass stage of at least an inch and a half in
diameter. A compound microscope is useful for all minute
investigation, and is essential in the study of vegetable anatomy
and of all lower cryptogamic botany.

772. For making thin slices, a razor is the best knife ; for dis-
section on the stage of the simple microscope, beside needles,
small scalpels or some of the cutting instruments used by ocu-
lists are very convenient. But an expert hand is able to do
almost every thing with a common knife or scalpél and a pair
of mounted needles. Slender forceps are almost indispensable :
those made for the use of dentists are the best.

773. Analysis. In the examination of an unknown plant with
a view to its determination, its whole structure should be made
out, so far as the materials allow, before a step is taken to
ascertain its name and place in the system. In respect to the
stem, its duration and consistence and its internal structure,
whether exogenous or endogenous, are to be noted. As to the
foliage, the venation and the phyllotaxy, also the presence or
absence of stipules, are most important. The anthotaxy or
inflorescence is to be examined and referred to its proper type.
In the flower, the numerical plan and symmetry, its ground-plan
and the nature of the deviations from the general or the family
type, are to be considered; also the estivation or arrangement
of the parts in the bud, the character and extent of coalescence
and adnation ; the manner in which the anther is borne upon the
filament, and its place and mode of dehiscence, &c. _ Note also
whether, when the blossom is hermaphrodite, the anthers and the
stigmas mature at the same or at different periods. The placen-
tation and the character and position of the ovules should be

SPECIMENS. atk

determined. Two sections of the flower should be made: one of
them vertical and directly through the centre, in the manner of
Fig. 336-341, —this will display the adnation, insertion, &c.,
of all the parts; the other transverse and through the middle of
the 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 zestivation. (Fig. 351, 886, 398,
&c.) Not rarely fruit and seeds are to be had at the same time,
or upon the same specimen, and these are equally to be investi-
gated. In fresh seeds, even those of minute size, the embryo
may almost always be extracted or brought to view under the
microscope, either by tearing away the seed-coat with needles or
by sections with a keen knife. When hard and dry, they have
only to be soaked or slightly boiled.

774. Diagrams and also sketches of the parts should be made,
such as those referred to in the foregoing paragraph. Such
diagrams can be drawn by any one with a little practice ; and
they may be made to express the whole floral structure, even to
the coalescence and adnation.' But in the process of determina-
tion the student should beware of trusting wholly to his diagrams
and sketches without direct verification.

775. Dried specimens, when well prepared and in sufficient
abundance, in the hands of a skilled botanist are in most cases
but little inferior to fresh ones. When needed, flowers, or clus-
ters of blossoms, or fruits may be detached and prepared for
examination and dissection by somewhat prolonged soaking in
warm water or by a short immersion in boiling water. This re-
stores flower-buds and small flowers and fruits, or their parts, to
a condition not essentially unlike the living state. Consequently,
the Herbarium or Hortus siccus of the botanist is to him more
essential than the botanical garden, important as that may be.

776. Herborizing.? The collector’s outfit will essentially con-
sist of a Vaseulum or botanical box, a Portfolio, a Trowel, a pocket
Lens, and a small but stoutly covered Note-book. Some use a
portfolio only, others the botanical box ; but on a long excursion
it is well to have both. The former is preferable in most cases,
except when specimens are collected for the immediate use of a

1 See Eichler’s Bliithendiagramme (Leipsic, 1872, 1878), an admirable
work, which may serve as a model.

2 These articles, from paragraph 776 to 802 inclusive, were obligingly pre-
pared, at the author’s request, by Lyman H. Hoysrapt, 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.

at2 PHYTOGRAPHY.

ciass. When well stocked with paper, it is of almost unlimited
capacity ; and most plants of delicate texture (as many of the
smaller aquatics, and those with fugacious or delicate corollas)
need to be consigned directly to the paper in which they are to
be pressed, and to be kept meanwhile under some pressure.

777. The Vaseulum is very useful for holding plants that are
to be examined fresh, and for thick roots, large fruits, &ce. It
is made of tin, and should be of oyal-cylindrical shape, about 17
inches long and 4 by 6 inches wide, and provided with a light
strap to throw over the shoulder. The lid opens nearly the
whole length of one of the flat sides (15 by 44 inches, with one
fourth inch lap), is made to fit as close as possible, and fastens
by a simple spring catch. When no portfolio is used, a larger box
may be required. Plants may be kept fresh in such a box for
many days. Fora several-days excursion, when it is desirable to
bring home a large number of fresh plants, a tin chest, made some-
what after the pattern of an old-fashioned trunk, will be found
very convenient. It should be about 21 inches long, 10 inches
wide, and 10 inches high to the top of the convex and hinged
lid, which forms the whole top, and to which a handle is fitted.

778. A good form of Portfolio is made of two pieces of binder’s
board covered with enamel cloth, and fastening together with two
long straps with buckles. Handles similar to those on a carpet-
bag may be attached for carrying. The usual size of portfolio
is 18 by 12 inches, but 16} by 11} inches may be better, as
there would then be little danger of making specimens of too
great length for the herbarium. (784.) Or the back may be of
soft leather, an inch or so in width, and a light strap and buckle
at the front edge and at each end. The portfolio should contain
a good quantity of folded sheets of thin unsized paper, similar
to grocer’s tea-paper, and of a size only a little smaller than the
sides of the portfolio. Very thin manilla paper, or what is so
called, is exceilent for this purpose, being sufficiently bibulous
and rather strong.

779. The specimens as soon as gathered should be laid neatly
in these folded sheets (called specimen sheets), and kept under a
moderate pressure in the portfolio. The sheets with the spec-
imens are afterwards transferred to the home press, but the
specimens should be left continuously in their sheets through all
the changing of driers, until cured. Indeed, the specimens may
well remain in the sheets after drying, until wanted for mounting
or for exchanging. For fine specimens, the use of this specimen
paper is very important. Many plants are so extremely delicate
and sensitive that they will not bear the least handling without

HERBORIZATION. 373

curling and shrivelling, unless thus enclosed: also without these
sheets much time is lost in transferring small specimens one by
one from one drier to another in the drying process.

780. For digging up roots, bulbs, &c., a small and sharp
pointed triangular Zrowel or stout knife will answer. One of
the best ‘‘ diggers” is made from a large file. Let a black-
smith bend the lower half of the blade to a gentle curve, so that
the point will be about an inch out of the true line. Grind off
the teeth and re-temper the blade. The total length with
handle, which is over one third, should be about twelve inches.
A leather case may be made for convenience of carriage. The
advantages of this strong tool are many.’

781. A Note-book should be carried upon every excursion, in
which the station of rare plants, dates, colors, and various par-
ticulars which cannot be learned from the specimens, may
be recorded on the spot, instead of being left to uncertain
memory.

782. For most plants, the best time for collecting flowering
specimens is in the morning, soon after the dew has disappeared.
Vespertine flowers have to be secured earlier, or at nightfall.

783. Care should be taken to have the specimen of the proper
size, neither too small nor too large, and to comprise all that is
necessary for complete botanical illustration, — flowers, fruit,
and leaves, both cauline and radical when possible. Inex-
perienced botanists suppose that a small sprig, containing a
flower or two with a few leaves, will answer all purposes as a
botanical specimen ; but later he comes to know better, and also
learns that the flower is only one of the component parts of a
specimen, and not always the most important one. In various
genera and orders, the fruit is the most distinguishing character-
istic, as with the Potamogetons, the Cruciferze, the Umbellifere,
and the Cyperaceze. 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. Consequently, all the
organs have their value in an herbarium specimen, and each and
all should receive due consideration from the botanist when col-
lecting. Specimens may be often secured that exhibit both

1 [There is an English herborizing trowel of excellent quality, with blade
six or eight inches long, less than two inches wide, the sides slightly in-
curved, the stout shank an inch and a quarter wide, and one sixth of an
inch thick: this forms the whole back of the handle, the front of which is
a piece of lignum vite riveted fast to the steel. It is nearly impossible to
break it.]

374 PHYTOGRAPHY.

flowers and fruit in the same plant, or fruit may be frequently
obtained from more advanced plants at the same time. If not,
fruit must be collected later, as in case of shrubs and trees, of
which generally only a branchlet with flowers, or with flowers
and leaves, can be gathered first. But subsequently the fruit
and mature leaves, should always be taken, if practicable, from
the same individual as the flowers. Of dicecious shrubs or
trees, like the Willows, each species should be represented by
four pieces: first, the sterile and fertile catkins will have to be
obtained, and the respective individuals marked, so that later
corresponding twigs with mature leaves, stipules, and fruit may
be gathered, and the specimens rightly matched.

784. A specimen should be so arranged as to be no larger
when pressed than can be neatly mounted on the herbarium
paper. A slender plant not over three feet in height should
generally be preserved entire, root and all. This can be done
by bending or partially breaking it at one, two, or three places,
and doubling so that the sections will not rest upon each other
in drying. If broken twice, it may be neatly arranged in the N
form when put in portfolio. Very large herbaceous plants will
have to be divided and the parts preserved separately, or, better,
take a suitable portion of the upper stem, having leaves, flowers,
and fruit, and a convenient part of the lower stem containing
radical leaves and with it sufficient root to show whether the
plant is an annual, biennial, or perennial. Thick stems, roots,
tubers, bulbs, and the like, should be divided or thinned down
with a knife, but in such a manner that the original shape can
be easily made out.

785. Carices should be always collected when the fruit is full-
grown, but not so ripe as to fall away. So also should other
Cyperacee ; yet it is well to collect also earlier specimens of
these in flower. Grasses, on the other hand, should generally
be collected soon after they come into blossom. For when
mature the spikelets in many species break up and fall away in
drying. The culm, leaves, and root of Sedges and Grasses
should be preserved, as well as the inflorescence. The root is
no less important. Cespitose species should be so collected and
preserved as to show the tufted character. The culms of most
sedges and grasses act stubbornly when bent for arrangement
in portfolio or press, and are not disposed to stay in place. This
difficulty is promptly remedied by crushing with the teeth the
angles made by the bending. Or these may be thrust through
slits of paper. In drying Sedges and Grasses, very moderate
pressure should be employed.

HERBORIZATION. 3875

786. Some aquatic plants (Algz especially) are so soft and
flaccid that, to secure them in their proper shape, they must be
placed in clear water and floated out by inserting beneath them
the paper on which they are to remain permanently, cither the
regular mounting paper, or a thinner white paper which when
dry can be pasted on the herbarium sheet. If likely to adhere
to the sheet or drier above them in the press, a piece of oiled or
stearine paper may be laid directly on the specimens to prevent
their sticking. Also viscous or glutinous plants which are liable
to adhere to the sheets enclosing them may be sprinkled with
Lycopodium spores, powdered soapstone, or some similar sub-
stance.

787. The name of the plant if known, but by all means the
locality and date of collection, with any other descriptive re-
marks regarded necessary, should be written on a ticket or on
the sheet when it is put into the press. Never omit to record
the time and place of collection, as a specimen of unknown date
and locality loses much of its value and interest.

788. Drying Specimens. ‘The chief requisite for good herba-
rium specimens is the extraction of the moisture from the green
plant as rapidly as possible under a pressure which obviates
brittleness. This is to be affected by placing the thin sheets
containing the specimens between layers of bibulous paper, called
driers, and applying moderately strong pressure to the pile.
For driers nothing can be better than thick blotting paper,
except that it is too expensive, and the same may be said of
an English drying paper made for the purpose. Equally good
driers are made of the thick and felt-like brown paper which,
after saturation with coal-tar, is here largely used under the
clapboards of wooden houses and under slate-roofing. It is a
cheap material, and is to be obtained, cut into sheets of 18 by
12 inches. Or driers may be made of old newspapers or of any
soft wrapping paper, cut or folded to the proper size, and
stitched (very expeditiously by a sewing-machine), or joined by
eyelet paper-fasteners at two corners, in packages of a dozen or
more leaves to a drier. It is well to have a large supply of driers
and specimen-sheets ready for use.

789. A half dozen or more pieces of thin boards, 18 inches
long and 12 inches wide, should be provided. They are used
at the top and the bottom of the pile when pressing, and also
for dividing it into suitable sections, especially for separating
the packages of plants which were put into press at different
periods, and dividing up these packages themselves, if too large.
For the plants dry better in small sections and with the pressure

310 PHYTOGRAPHY.

evenly distributed. Hence it is best to have these sections not

over five or six inches in thickness, nor should the pile itself .
be carried too high, never exceeding two feet. Painted binders’

boards may be used, instead of the common boards, to separate

interior divisions. Some botanists use a kind of lattice made of

two layers of thin strips or laths, crossing each other. This is

said to allow free escape of the moisture by evaporation, and so

to accelerate drying, as in the case of the wire press.

790. For giving pressure, various ways haye been contrived.
The Serew-press is convenient and compact, but objectionable,
because it does not follow up the pressure as the plants shrink
in drying. This objection does not apply to the Lever-presses, but
they are usually unwieldy. Fortunately, one of the best forms
of the drying-press, as well as the simplest and cheapest, is
merely a board with weights placed on the top of the pile of
specimens. Here the pressure is continuous, constantly follow-
ing the shrinkage of the plants. The weight ona pile should
vary from 25 to 100 pounds, according to the nature of the
specimens and the quantity in the press. On an average, 60
pounds is sufficient for most plants. If much greater pressure
is used, there is danger of crushing the more delicate parts of the
specimen, and thereby impairing its scientific value. For weights,
bars or masses of iron may be used, boxes filled with sand,
stones, and the like.

791. Specimens brought home in the botanical box must be
placed in such thin specimen-sheets as are used in portfolio.
In putting plants in specimen-sheets, whether in portfolio or
press, it is well to take some pains to spread out the specimens
neatly ; for a little care now may save much later labor. How-
ever, with most species, any carelessness in this respect can
be remedied at the first change of driers. But there are some
plants, previously referred to, so peculiarly sensitive that what-
ever adjustment they receive must be given at the time they are
first placed in their sheets.

792. Aithough plants can, if necessary, be kept fresh for several
days in box or portfolio, on returning from a collecting trip they
should be transferred to the home press as early as possible. In
the transference, particular care should be taken to straighten
out and remove all folds and crumpling of the leaves, petals,
fronds, &c., and to arrange the specimen as naturally as possi-
ble, so as to show the proper habit. Both sides of the flowers
and leaves should be exhibited. Plants that were put directly
into press should receive this special attention at the first
change of driers. which on this account should be made within

» Ti

HERBORIZATION. 377
several hours afterwards. The stubbornness and elasticity, so
troublesome in specimens when first put in, will then have
mostly disappeared, and the whole specimen will be found suffi-
ciently flaccid to have every part stay as arranged. If this first
change is deferred longer than ten or twelve hours, the speci-
mens of many species become too dry for making the alterations
required. At this time small pieces of bibulous paper may be
placed between leaves, or other portions of the plant which over-
lap, to prevent moulding or discoloration, and to hasten drying.
It is well to change these fragments of paper with the driers for the
first day or two: afterwards they may remain with no detriment.

793. To have the specimens retain their natural color and
general appearance, they should be dried as rapidly as possible ;
and this result is best secured by frequent changing of the driers.
These should be changed at least once a day for the first four or
five days, and afterwards every 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 expose them to the sunshine, and bringing them in
hot to make the change at once, or as soon as possible.

795. The number of driers interposed between the specimen-
sheets should depend upon the plants and the frequency of the
changes: two will suffice when the driers are changed very often ;
but more must be employed when the plants are thick and succu-
lent. Uniform pressure 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 about some of the larger flower-heads of the Composite, &e.

796. The time required to dry specimens varies with different
species and with the season: it depends also on the frequency
of the changes and the temperature of the driers. By changing
daily, the time is usually from four or five days to a week. But,
with two changes a day for the first day or two and with heated
driers, the process may be completed in half the usual time, and

3878 PHYTOGRAPHY.

the specimens will be in much finer condition. An experienced
collector has no difficulty in ascertaining whether a plant is com-
pletely cured or not, while to a novice it is often a matter of
uncertainty. A thoroughly dried plant can be usually told by
its peculiar hay-like rattle when disturbed; also by placing the
plant against the cheek. If there is a sensation of coolness, the
plant is still moist.

797. If the thick leaves of fleshy plants are immersed for
a few moments in hot water, the period of desiccation will be
greatly hastened; but they frequently turn dark as a conse-
quence of the immersion. The drying of such plants, and
particularly of the Monocotyledons, may be advantageously ex-
pedited by placing them between several driers and ironing them
with hot irons. Small plants may be very neatly dried in old
books. Very beautiful specimens may be made by placing the
plant in a tall and narrow vessel, and pouring over it a sufficient
quantity of clean and dry sand. When the moisture is absorbed,
it may be flattened in a press.

798. In shifting the driers of a collection, place the package
to be changed at the left hand on the table or counter, the new
pile in front with its length parallel to the person, — a position
the most favorable for giving any needed attention in arranging
specimens, — while fresh driers may be placed at the right hand,
or beyond the pile in front. Thus arranged, the sheets of speci-
mens can be rapidly shifted into their fresh driers.

799. The moist driers may be spread out in the sunshine to
dry, or strung on a line in a warm room, or in the open air, if
not too windy. Very moist driers may be thoroughly dried
within an hour, if spread in the hot sunshine. In inclement
weather, they must be dried by the fire.

800. To recapitulate the most important points in good speci-
men-making: Use specimen-sheets to hold the plants undis-
turbed during the whole process of drying: use plenty of the
most bibulous driers, sun-dried and heated when practicable: do
not make the piles too large: make the first shift of driers within
a few hours, at that time making all needed adjustment of the
flaccid specimens: change the driers twice a day for the first
day or two.

801. For collecting and preserving specimens on a journey,
or when moving from place to place, some modification of the
stationary press is requisite. The Z’ravelling-press nmust be porta-
ble: accordingly the pressure is applied by strong leather straps
with buckles. There should be three straps, one girding the
package around each end, and one lengthwise. The top and

HERBORIZATION. 379

bottom, if of thin boards, must be cleated, or compounded of
wood with the grain in opposite directions, or very stout binder’s
board or trunk-board may be advantageously used. This should
be covered with coarse cotton or linen cloth, glued fast and well
painted. While stationary, the pressure may be given by means
of weights when more convenient.

802. The Wire press, now much in use, is a press of this porta-
‘ble kind, in which the boards are replaced by sheets of wire net-
ting, with wide meshes, and surrounded and strengthened by a
strong but light iron border. Straps with buckles are used to
hold the parts’ and contents together and to apply the pressure,
as in the ordinary trayelling-press. Besides its portability, the
advantages of such wire presses are that, in a small way, they
may serve both as portfolio for collecting and as press for dry-
ing; also that, as the drying takes place mainly by evaporation
instead of absorption, much less paper is required, and the trouble
of changing the driers is saved.' In fair weather, the press filled
with plants may be hung in the wind or sunshine, in foul weather
near a fire. The disadvantage is that specimens dried in this
way are apt to be brittle. To use this system advantageously,
the botanist should have at least two such presses in operation,
one for collecting, while the other is in use for, drying.

803. Poisoning. Dried specimens are liable to the depreda-
tions of certain insects, especially of their larvee. The principal
pest is a smail brown beetle, Anobium paniceum, L.; the perfect
insect does considerable damage, the larva vastly more. Plants
with milky juice, such as Asclepiadezx, Apocynace, and Eu-
phorbiacez, those containing bitter principles, such as Gentians
and Willows, and generally such plants or such organs as con-
tain much protoplasm or azotized matter, are most subject to
attack. Ranunculacee, Umbelliferze, and Composite are seldom
spared ; while Labiatez mostly escape, probably on account of the
volatile oil which they contain. Even Ferns are liable to have the
parts of fructification eaten away. To a certain extent, the im-
pregnation of the herbarium-cases with camphor, naphthaline,
or strong-scented oils, may exclude the vermin. But safety is
secured only by poisoning.

804. The proper poison is corrosive sublimate, dissolved in
strong (95 percent ) alcohol. Drop into the alcohol as much cor-

1 Prof. A. Wood seems to have been the first to call the attention of
American botanists to this system, which he has earnestly advocated.

An improved form of this wire press, well adapted both for collecting
and pressing in moderate quantity, is made and sold, at a small price, by
Paul Ressler, optician, at New Haven, Connecticut.

380 PHYTOGRAPHY.

rosive sublimate as it will take up, then add a trifle more of alcohol,
so as to keep the solution just below the point of saturation.
The stronger the solution the better, except that, at full satura-
tion and where copiously used, an efflorescence may sometimes be
left on the surface of the poisoned specimens upon the evapora-
tion of the alcohol. Some add to the solution some carbolic
acid, at the rate of a fluid ounce to each quart of alcohol. The
solution may be applied with a soft brush (one with no metal in
its fastening), or by a dropping bottle, or even the specimens
may be dipped in the solution placed in a flat porcelain dish.
The brush (using a pretty large and soft one) is the most con-
venient and efficient. The moistened specimens should be placed
between driers and in shallow piles until the alcohol evaporates.

805. Thoroughly poison all specimens before admitting them
to the herbarium. It is well to poison all specimens whatever,
as soon as they are made or at the close of the botanizing sea-
son, as well those intended for exchanges as for the collector’s
own herbarium.

806. Keep all specimens between sheets of paper, or within
folded sheets, not too crowded or overlaid, away from dust, and
in a perfectly dry place, so as to avoid mould. When attacked
by mould, the corrosive-sublimate solution should be applied.
A properly dried specimen, duly cared for, should be as lasting
as the paper which holds it.

807. The Herbarium, called by the earlier botanists Hortus
Siccus, is a collection of dried specimens, named and systemat-
ically arranged. It is indispensable to the working systematic
botanist, and every devotee of botany should possess, or have
access to an herbarium containing representatives of the plants
of the immediate vicinity or district, if not of the whole country.
Or an herbarium may be restricted to a particular family of
plants, made the object of special study. A general herbarium
should contain specimens representing all the natural orders and
as many of their genera and species as possible.

808. The form of the herbarium as to the size of its sheets is
considerably variable. That of Linnzeus is of the size of foolscap
paper: this would now be universally regarded as much too
small. The principal British herbaria adopt the size of 163 by
10} inches, which is rather too narrow, rarely permitting two
specimens of the same species of any considerable size to be
placed side by side on the same sheet. In the United States,
16} inches in length by 11? in width is adopted; that is, for the
genus-covers, the species-paper being a quarter of an inch nar-
rower.

THE HERBARIUM. 381

809. The specimens representing each species may either be
laid within a doubled sheet, loosely (as in some European her-
baria), or fastened in place by narrow slips of gummed paper
(which is much better), or else they may be 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 any time be
exchanged for better ones. But a large herbarium on this plan
becomes cumbrous and inconvenient for ready reference and
comparison.

811. The best plan in a large herbarium, and one much to be
consulted, is to attach the specimens completely, by any kind
of strong and light-colored glue, to single sheets, or rather half
sheets. The specimens are thus safe from injury under reason-
able handling, and can be turned over and examined with as
much facility as a series of maps or engravings. The species-
paper should be of writing-paper stock, or of equal firmness, of
compact texture, well sized and calendered, and of a weight in
size of 164 by 11} inches of about 18 pounds to the ream of
480 flat sheets. The paper should be furnished square-cut on
all sides, in the manner of ** flat cap.” Stiffness is the great
desideratum.

812. In no case should more than one species be knowingly
attached to the same sheet. But of very many species there will
be room for more than one specimen. And specimens from dif-
ferent localities, of different forms, and in various stages of
flowering and fructification, are always desirable. The full name
of the plant should be written at the lower right-hand corner of
the sheet, or a ticket should there be attached by glue or traga-
canth paste. Each specimen should have its ticket, similarly
attached, or a memorandum upon the sheet, indicating the hab-
itat or the special locality, date of collection, name of collector,
and any other desirable information which the specimens them-
selves do not furnish. When there are loose flowers or fruits, or
when any of these have been detached for dissection and micro-
scopical investigation, it is well to preserve them, placing them
in little paper pockets or envelopes and pasting these upon the
sheet close to the specimen to which they pertain. Sketches of
parts dissected may be drawn upon the sheet. Notes and mem-
oranda received with the specimen or too extended to be entered
upon the sheet may be folded, inserted in such envelopes, and
made fast to the sheet. Many botanical collections are distrib-
uted with printed tickets. These, and ail authenticating tickets

382 PHYTOGRAPHY.

or notes, should be attached to the sheet near to the specimen
they belong to. In view of this, printed and written tickets
should be of small size.! A ticket which exceeds four by two

1 All printing on an herbarium ticket should be in plain type; and fancy
borders, uselessly occupying room, should be avoided. If any border is
thought needful, it should be of plain lines. Itis not desirable to parcel out
the space on a ticket with separate lines and headings for habitat, date of
eollection, time of flowering or fruiting, name of collector, and the like.
These particulars may conveniently be entered at the bottom or top of the
ticket,as may be convenient, leaving the rest of the space free for the name
of the plant, the authority, and perhaps a synonym.

Tickets for specimens distributed among other botanists may well have a
head-line indicating the source, such as “ EX COLL. C. C. PARRY,” or “ EX HERB.”
or, in English, ‘“‘ From roe Herparictm or” the botanist who communicates
the specimen. The following may serve as an example of a simple ticket
for the sending out of dried specimens, and of the way in which the ticket
may be filled out with the name of the plant, its habitat and station, name
of collector and time of collection

EX HERB. A. GRAY.

Sh qprke gia caruka, Tarr.

tae ae P ar? , le Gee aoe Joe f-¢. aad,
af J. Jr YAS rag, Aly, 1877.

For the botanist’s own herbarium, it is well to use a blank ticket with a
printed heading like the specimen above, but with the “ex ” omitted.

When a considerable collection is made in any particular botanical explo-
ration or excursion, and numerous or several specimens of the same species
are gathered, to be distributed among botanists in the way of exchange or
otherwise, these are commonly given out under numbers and with a printed
heading to a special ticket. The following is an approved form of such a
ticket, and of the mode in which it may be filled up in writing by inserting
the name of the species, the locality, &e.

ALPINE FLORA OF THE ROCKY MOUNTAINS.
Coll. C.C. PARRY. 1872.

No. IVF

CoLorabDo, trae 6 Cee
Geir ove t alt. 2 hip.

THE HERBARIUM. 383

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 are
most commodious.

813. The sheets of ali the species of the same genus, when
not too numerous, or of a particular section of it, or any conyven-
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 164 by 113 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 1? 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 corner ; that is, on the upper face next
the back: ator near the lower right-hand corner, the name of the
contained species may be written either with a pencil or in ink.

814. The genera should be arranged in the herbarium accord-
ing to some systematic work, and numbered accordingly on the
covers.

815. The herbarium must be preserved in close cabinets or
cases free from the access of dust. Tin cases, just deep and
wide enough to receive comfortably the genus-covers, and about
six inches high, the hinged lid being one end, may be recom-
mended for a small collection, as they are dust and _ insect
proof, are portable, and may readily be arranged on shelves.
But, for any herbarium of considerable size and continued growth,
wooden cabinets with well-fitted doors are to be preferred; the
interior of the cabinets being divided into pigeon-hole compart-
ments, fully 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 board, or upon a flap of card-board affixed to it.
Moreover, it is well to write the name of the order upon each
genus-cover.

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

384 PHYTOGRA PHY.

of compartments. Such cabinets can be increased in number as
required, are portable, and can be disposed in any order, side by
side or one surmounting another, as may be most convenient.
The doors should be so constructed as to open and shut readily,
but to close tightly, so as to exclude dust and insects.?

1 An excellent plan for small and inexpensive herbarium cabinets, of a
portable character, is proposed and illustrated by Dr. Parry, in the American
Naturalist, 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 compartments high in Dr. Parry’s plan, but double the number
might be better): the entire front consists of a pair of doors meeting in the
centre, there fastened by a flush spring catch; the doors bevelled on the inside,
with a corresponding bevel on the case, to which they are attached by out-
side hinges, so that in opening at a right angle there are no sharp corners to
hinder the drawing out of the herbarium papers; also allowing the cases
to stand close side by side, as well as one upon another, without interfering
with the free opening of the doors. These, moreover, may swing quite back
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.

ABBREVIATIONS.

i. OF NAMES OF BOTANISTS AND BOTANICAL AUTHORS.

Ach.
Adains.

Afz.

All.
Amm.
Anders.
Andr.
Andrz.
Aresch.
Arn.
Arrh,
Asch.
Aubl.

Bab.
Bail.
Balb.
Baldw.
Balf.
Barn.
Barr.
Bart.

W Bart.
Bartr.
Bartr. f.
Bauh.
Beauv.
Benj.
Benn.

A. Benn.
Benth.
Berg.
Berk.

Acharius.
Adanson.
Afzelius.
Agardh.

C. A. Agardh.

J. G. Agardh, son.

Aiton.
Allioni.
Amman.

Andersson of Stockholm.

Andrews.
Andrzejowski.
Areschoug.
Arnott.
Arrhenius.
Ascherson.
Aublet.

Babington.

Baillon.

Balbis.

Baldwin.

Balfour.

Barnéoud.

Barrelier.

Benj. Smith Barton.

W™- P.C. Barton, nephew.

John Bartram.

Wm. Bartram.
Bauhin.

Palisot de Beauvois.
Benjamin.

J J. Bennett.

A. W. Bennett.
Bentham.

Bergius.

M. J. Berkeley,

Berkh. = Berkhey.

Berken. Berkenhout.

Berland. Berlandier.

Bernh. Bernhardi.

Bert. Bertero.

Bertol. Bertoloni.

Bess. Besser.

Bieb. Marschall von Bieberstein.

Bigel. Jacob Bigelow.

Bisch. Bischoff.

Behm. Boehmer.

Boerh. Boerhaave.

Boiss. Boissier.

Boland. Bolander.

Bong. Bongard.

Bonpl. Bonpland.

Bork. Borkhausen.

Borsz. Borszcow.

Brack. Wm. D. Brackenridge.

Brebis. Brebisson.

Bref. Brefeld.

Brew. &§ | W. H. Brewer & Sereno
Wats. Watson.

Brid. Bridel.

Brong. Brongniart.

Brot. Brotero.

Brouss. Broussonet.

Br., R. Br. Robert Brown.

PP) Br: Patrick Browne.

Brunf. Brunfels.

Buckl. Buckley.

Bull. Bulliard.

Burm. Burman.

Buzb. Buxbaum.

Cam. Camerarius.

Camb.

Ganken: t Cambessedes.

386 ABBREVIATIONS.

Campd. = Campdera. Eat. = Amos Eaton.
Cand. DeCandolle, usually DC. D.C. Eat. D.C. Eaton, grandson.
Casp. Caspary. Edgew. Edgeworth.
Cass. Cassini. Edw. Edwards.
Catesb. Catesby. Ehren. Ehrenberg.
Cav. Cavanilles. Ehirh. Ehbrhart.
Cerv. Cervantes. Eichl. Kichler.
Cham Chamisso. Hiseng. Kisengrein.
Chapm. A. W. Chapman. Ell. Elliott.
Chav. Chavannes. Endl. Endlicher.
Chois. Choisy. Engelm. Engelmann.
Clayt. Clayton. Engl. Engler.
Clus. Clusius. Eschs. Eschscholtz.
Collad. Colladon. Eschw. Eschweiler.
Colm. -Colmeiro. Ettingsh. Ettingshausen.
Comm. Commelin.
Corn. Cornuti. Fendl. Fendler.
Coss. Cosson. Feuil. Feuillée.
Cunn. Cunningham, A. or J. Fingerh. ¥ingerhuth.
Curt. Wim. Curtis. Fisch. Fischer.
M.A.Curt. M. A. Curtis. Forsk. Forskal.
Forst. Forster.
Dalech. Dalechamps. Fourn. Tournier.
Dalib. Dalibard. Frresen. Fresenius.
Darl. Darlington. Freye. Freycinet.
DC. Freel. Freelich.

{ A. P. DeCandolle.

A.DC. Alphonse DeCandolle,son. | Goartn. J. Gertner.
Cas. DC. Casimir DeCandolle, the | Gertn. fC. 'T. Geertner.

Decne. Decaisne. [grandson. | Gardn. Gardner.
Deless. Delessert. Garid. Garidel.
Dennst. Dennstedt. Gasp. Gasparrini.
Desc. Descourtilz. Gaud. Gaudin.
Desf. Desfontaines. Gaudich. Gaudichaud.
Desj. Desjardins. Germ. Germain.
Desmar. Desmazieres. Gesn. Gesner.
Desmoul. Desmoulins. Gilib. Gilibert.
Desv. Desvaux. Ging. Gingins de Lassaraz.
Dicks. Dickson. Gis. Giseke.
Diesb. Diesbach. Gled. Gleditsch.
Dieter. Dieterich. Gleich. Gleichen.
Dietr. Dietrich. Glox. Gloxin.

Dill. Dillenius. Gmel. J. G. Gmelin.
Dillw. Dillwyn. C. Gmel. C.C. Gmelin of Baden.
Dod. Dodonzus (Dodoens). S. Gmel. S. G. Gmelin.
D’ Orb. D’Orbigny. Godr. Godron.
Dorst. Dorstenius. Gepp. Geeppert.
Dougl. Douglas. Good. Goodenough.
Dre}. Drejer. Gren. Grenier.
Dryand. Dryander. Grev. Greville.
Dufr. Dufresne. Griseb. Grisebach.
Duham. Duhamel du Monceau. Gren. Greenland.
Dumort. | Dumortier. Gron. N

Gronovius.
Dun. Dunal. Gronov. $

ABBREYIATIONS.
Guett. == Guettard. Jacq. =
Guib. Guibord. Jacq. f:
Guillem. |Guillemin. J. St. Hil.
Guimp. Guimpel. Jord.
Gunn. Gunnerus. Jungh.
Guss. Gussone. Juss.
Adr. Juss.
Hagenb. Hagenbach.
Fall. Haller. Kemp.
Ham. Hamilton. iarct:
Llanb. Hanbury. Kaulf
Haust. Hanstein. sents
Havtin. Hartmann. oe ;
irschl.
Hartw. Hartweg. Kit
Harv. Harvey. K es
alr.
Hass. Hassall. Korth
Flassk. Hasskarl. jon ;
ostel.
Hausm. Hausmann. Kremp
Haw. Haworth. Ve 4
. romo.
Hebens. Hebenstreit. fee
Hedw. Hedwig. ;
Hegelm. Hegeliaier.
Hegetsch. egetschweiler. L.
Heist. Heister. Labill.
Fleldr. Heldreich. Lest.
Helw. Helwing. Lag.
Hemsl. Hemsley. Lall.
Flenck. Henckel. Lam.
FHlenfr. Henfrey. Lamb.
FHensl. Henslow. Lamour.
Herb. Herbert. Langsd.
Herm. Hermann. La Peyr.
Hild. Hildebrand. La Pyl.
TTochst. Hochstetter. Ledeb.
Ho fjin. G. F. Hotfmann. Lehin.
H1. Hoff. Hermann Hoffmann Lem.
Hoffimanns. Hoffmannsegg. Lesq.
FHlofin. Hofmeister Less.
Hohen. Hohenacker. Lestib.
Holmsk. | Holmskiold. Lév.
Flomb. Hombron. LD Her.
Fook. Wm. J. Hooker. LT Herm.
Hool:. f. J. DY. Hooker, son. Liebm.
Hopk. Hopkirk. Lightf:
Hornem. Hornemann. Lili).
Flornsch. Wornschuch. Lindb.
Florsf. Horsfield. Lindbl.
Foust. Houston. Lindenb.
Houtt. Houttuyn. Lindh.
Huds. Hudson. Lindl.
Hueb. Huebener. Linn.
Hunb. Humboldt. Linn. f.
HBK. 3 Humboldt, Bonpland, and | Lodd.

1 Kunth.

Leff.

337

N. J. Jacquin.

J. F. Jacquin, son.
Jaume St. Hilaire.
Jordan.

Junghuhn.

A. L. Jussieu.
Adrien Jussieu, son.

Kempfer.
Karsten.
Kaulfuss.
Kindberg.
Kirschleger.
Kitaibel.
Keireuter.
Korthals
Kosteletzky.
Krempelhuber.
Krombholz.
Kuetzing.

Linneus.

La Billardiere
Lestadius.
Lagasca.
Lallement. | Marck).
Lamarck (Monnet de La
Lambert.
Lamouroux.
Langsdorf.

La Peyrouse.
La Pylaie.
Ledebour.
Lehmann.
Lemaire.
Lesquereux.
Lessing.
Lestiboudois.
Léveille.

L’ Heritier.

L’ Herminier.
Liebmann.
Lightfoot.
Lilijeblad.
Lindberg.
Lindblom.
Lindenberg.
Lindheimer.
Lindley.
Linneus. Also Z.
C. Linnzus, son.
Loddiges.
Leefling.

388 ABBREVIATIONS.
Les. = Leselius. Naud. =
Los. Loiscleur-Delongschamps. | Neck.
Loud. Loudon. Nees or !
Lour. Loureiro. N.ab E.
Ludw. Ludwig. T. Nees
Lumn. Lumnitzer. Nestl.
Lyngb. Lyngbye. Newb.
Newm.
Macf. Macfadyen. Negg.
Macgil. MacGillivray. Nois.
Magn. Magnol. [stein. | Nord.
M. Bieb. Marschall von Bieber- | Jot.
Marsh. Humphrey Marshall. Nutt.
Mars. Marsili. Nyl.
Mart. Martius. Nym.
Mass. Massalongo.
Mast. Masters. (Ed.
Maxim. Maximowicz. (rst.
Med. Medikus or Medicus. Oliv.
aes Meisner or Meissner. Heut
Meneq. Meneghini. Orph.
Menz. Menzies. Ort.
Mert. Mertens. Oudem.
Metten. Mettenius.
Mich. Micheli. P.deBeauc.
eer André Michaux. - aa
Michx. f. ¥. A. Michaux, son. Park.
Midden. |Middendorff. Parl.
Mill. Philip Miller. Pasq.
Mill. J. John S. Mueller or Miller. | Pav.
Miq. Miguel. Perl.
Mirb. Mirbel. Pers.
Mitch. John Mitchell. Philib.
Mitt. Mitten. Planch.
Moe. Mocino. G. Planch.
Molk. Molkenboer. Pluk.
Mont. C. Montagne. Plum.
Moq. Moquin-Tandon. Peepp.
Moric. Moricand. Poir.
Moris. Morison. Poit.
Morr. Morren. Poll.
Moug. Mougeot. Post.
Muell. Arg. J. Mueller of Argau. Pourr.
F. Muell. Ferdinand Mueller. Pringsh.
O. Muell. Otto Muellerof Denmark. | Pritz.
Muhl. Muhlenberg. Putter.
Munt Munting.
Murr. J A. Murray. Rabenh.
A. Murr Andrew Murray. Radlk.
Raf:
Nace. Naccari. Rasp.
Neg. Negeli. Red.

Naudin.
Necker.
C. F. Nees von Esenbeck.

T.F.L. Nees von Esenbeck.
Nestler.
Newberry.
Newman.
Neeggerath.
Noisette.
Nordstedt.
Notaris.
Nuttall.
Nylander.
Nyman.

Eder.

(Ersted.

Olivier.

D. Oliver.

A. or C. d’Orbigny.
Orphanides.
Ortega.

Oudemans.

Palisot de Beauvois.
Pallas.

Panzer.
Parkinson.
Parlatore.
Pasquale.

Pavon.

Perleb.

Persoon.

Philibert.

J. E. Planchon.
Gustave Planchon.
Plukenet.

Plumier, Lat. Plumerius.
Peeppig.

Poiret.

Poiteau.

Pollich.

Postels.

Pourret.
Pringsheim.
Pritzel.

Putterlich.

Rabenhorst.
Radlkofer.
Rafinesque-Schmaltz.
Raspail.

Redoute.

ABBREVIATIONS.

Reich. = Reichard. Scop. =
Reichenb. WH. G. L. Reichenbach. Seem.
Reichenb. f. H. G. Reichenbach, son. Sendt.
Reinw. Reinwardt. Seneb.
Reiss. Reisseck. Ser.
Retz. Retzius. Seub.
Reut. Reuter. Sibth.
Rich. L. C. Richard. Sieb.
Rich. f. : : Sieb.
ac pad } Achille Richard. Rae
Liichards. John Richardson. Sow.
Richt. Richter. Spenn.
Ridd. Riddell. Spreng.
Riv. Rivinus. Sternb.
Reh. Rebling. Steud.
Rem. J. J. Roemer. Stev.
M. J. Rem.M. J. Roemer. Sull.
eg Remer & Schultes. iat

Sch.
Rep. Reper. Targ:
Rohrb. Rohrbach. Ten.
Rostk. Rostkovius. Thoms.
Rothr. Rothrock. Thuill.
Rottb. Rottbell. Thunb.
Rottl. Rottler. Thurb.
Roum. Roumegere. Thurm.
Roxb. Roxburgh. Tod.
[oy. Royen. Torr.
Rudb. Rudbeck. Torr. §
Rupr. Ruprecht. Tourn.

Tratt.

Sace. Saccardo. Traut.
Sadi. Sadler. Trev.
St. Hil. A. Saint-Hilaire. Trin.
Salish. Salisbury. Tuck.
Salm-Dyck.Prince Jos. Salm-Riffer- | Turcz.
Sauss. Saussure. [schied-Dyck. | Turn.
Schimp. | Schimper. Turp.
Schk. Schkuhr.
Schlecht. Schlechtendal. Vaill.
Schleich. Schleicher. Veill.
Schomb. Schomburgh. Vauch.
Schrad. Schrader. Vent.
Schreb. Schreber. Vill.
Schueb. Schuebeler. Vis.
Schult. Schultes. “ttad.
Schultz | C. H. Schultz, Bipontinus iv.

Bip. (Zweibrucken) Vog.
Schum. Schumacher.
Schnitzel. Schnitzlein. Wahl.
Schwagr. Schwegrichen. Wahist.
Schwein. Schweinitz. Walds.
Schweinf. Schweinfurth. Wall.
Schwend. Schwendener. Wall.

G

~

Scopoli.
Seemann.
Sendtner.
Senebier.
Seringe.
Seubert.
Sibthorp.
Sieber.
Siebold.
Solander.
Sowerby.
Spenner.
Sprengel.
Sternberg.
Steudel.
Steven.
Sullivant.
Swartz.

Targioni-Tozetti.

‘Tenore.

Thomas Thomson.
Thuillier.
Thunberg.
Thurber.
Thurman.

Todaro.

Torrey.

.Torrey & A. Gray.

Tournefort.
Trattinick.
Trautvetter.
Treviranus.
Trinius.
Tuckerman.
Turezaninow.
Turner.
Turpin.

Vaillant.

Veillard or Vieillard.
Vaucher.

Ventenat.

Villars, or Villar.
Visiani.

Vittadini.

Viviani.

T. Vogel.

Wahlenberg.
Wahlstedt.
Waldstein.
Wallich.
Wallman.

389°

“390

Wallr. = Wallroth.

Walp. Walpers.

Walt. Walter.

Wang. Wangenheim.
Warm. Warming.
Wats. P. W. Watson.

H.C.Wats.H. C. Watson.

S. Wats. Sereno Watson.
Web. W eber.

Wedd. W eddell.
Weinm. Weinmann.
Welw. Welwitsch.
Wender. Wenderoth.
Wendl. Wendland.
Wiks. Wikstrom.

ABBREVIATIONS.

Wildb. = Wildbrand.

Willd. Willdenow.
Willk. Willkomm.
Wils. Wilson.
Wimm. Wimmer.
Wisliz. Wislizenus.
With. Withering.
Woode. W oodville.
Wulf. Wulfen.
Zanard. Zanardini.
Zetterst. Zetterstedt.
Zuce. Zuccarini.
Zuccag. Zuccagini.

2. ABBREVIATIONS OF NAMES OF ORGANS AND TERMS
¢ USED IN BOTANICAL WRITINGS.

Est. Jéstate, in summer.

“Est. Estivation.

Alb. Albumen.

Anth. Anther.

Art. Artificial.

Auct., Auctt. Auctorum, of authors.

Aut. Autumnal.

B. or Beat. Beatus, “the late,” re-
cently deceased.

Br. Bract.

Cal. Calyx.

Cel. Celeberrimus, or Very cele-

brated.
Cent. Centimetre.
Cl. Clarissimus.
Char. Character.
Coll. Collection.
Cor. Corolla.
Cult. Cultivated.

Decim. or Dec™. Decimetre.

Deser. Description

Diff. Differentiz, the distinguishing
marks.

Ed. Edition.

Embr. Embryo.

Ess. Essential, as Char. Ess.

Excl. Excluding, or being excluded.

Excl. Syn. Excluding the synonym
or synonyms.

Fam. Family.

Fil. Filament of the stamen.

Fl. Flower (flos); Flora, or some-
times Floret, it flowers.

Fem. Female plant, flower, &c.

Fol. Folium, leaf.

Fr. Fruit.

Fructif. Fructification.

Gen. Genus or Generic.

Germ. Germen, Linnean name for
ovary ; also Germination.

H. Herbarium.

Hab. Habitat, place of growth;
sometimes for Habeo, I have.

Herb. Herbarium.

Hort. Hortus, garden.

Hortul. Hortulanorum, of the gar-
deners.

Ic. Icon, a plate or figure.

7/7. Mllustris, illustrious.

Ined. Unpublished.

Inf. Inferior.

Infl. Inflorescence.

Inv. Involucre.

Lat. Lateral, or relating to width.

Lin. Linea, a line (the 12th of an
inch).

Lit., Litt. In a letter or letters.

| 1. c. Loco citato, in the place cited.

ABBREVIATIONS. 391

Musc. Male plant, flower, &c. | Sep. Sepal.
Mill. or mm. Millimetre. Ser. Series.
Mss. Manuscripts. | Sice. Siccatus or Siccus, dried or dry.
Mus. Museum. Spec. or Sp. Species, or specimen.
N. or No. Number. | Spont. Spontaneous.
Nat. Natural. | Stam. Stamen or Staminate.
Nom. Nomen, name. Sup. Superior.
Obs. Observation. | Syn. Synonym or Synopsis.
Ord. Order. | T. or Lab. Tabula, plate.
Ov. Ovary. | ZY. Tomus, volume.
p. Page, or sometimes Part. V. Volume: sometimes for Vel, or;
Ped. Peduncle or Pedicel, or for sometimes Vide, see.
Pedalis, a foot long or high. : Var. Variety.

Peric. Pericarp.
Perig. Perigonium.

Pet.
Pist.

Place. Placenta.

Poll.
p- p-

Prodr. or Prod. Prodromus.

Rad.

Ram. Ramus, branch.
s. Seu, or Sive, Latin for or.

Veg. Vegetation, cliaracters of

Vern. Vernal.

v.s. Visa sicca, or Vidi siceam.

v.v. Visa viva, or Vidi vivam; the
first indicating that a dried speci-
men of the piant, the second that
the living plant has been exam-
ined.

» Ss. c. and v. &. &., indicates that
the dried specimen was cultivated
(c) or spontaneous (s).

Petal or Petiole.
Pistil.

Pollicaris, an inch long.
Pro parte, in part.

I

Radix, root; or Radical.

Sect. Section. v.v. c. and v. v. s., that the living
Segm. Segment. plant seen was cultivated (c) or
Sem. Semen, seed. spontaneous (s).

SIGNS.

1. SIGNS USED BY LINNAUS.

© _ An annual plant.
g A biennial.
2 A perennial.
5 A tree or shrub.
* Affixed to a reference, means that a good description will be found there.
+ Indicates an obscure or doubtful species.

‘)
@

2. SIGNS USED BY DECANDOLLE AND LATER
WRITERS.

A monocarpic plant, 7. e. which dies after once flowering and fruiting,
either annual or biennial, or of longer duration.

Annual.

Biennial.

Monocarpic perennial, such as Agave.

392 SIGNS.

Perennial herb.

Suffruter, an undershrub.

Frutex, a shrub.

Arbuscula, a tree-like shrub of ten to twenty-five feet in height.

Arbor, a tree.

A climbing plant.

An evergreen.

Male plant or flower.

Female plant or flower.

Hermaplirodite plant or flower.

Indefinitely numerous, e. g. 00-andra, polyandrous.

A sign of doubt. “ Thalictrum? Japonicum,’ doubts if the plant is
really a Thalictrum. “ Thalictrum Japonicum, Thunb.?” doubts if the
plant in hand is truly the species of Thunberg. Thalictrum Japoni-
cum, Thunb., Willd.? doubts whether Willdenow’s 7. Japonicum is
really that of Thunberg.

A sign of certainty. As “ Thalictrum anemonoides, Michx.! FI]. Bor. Am.
p. 822,” as used by DeCandolle, affirms that he nas seen an authentic
original specimen of this author. Affixed to the name of a collector,
as “ Virginia, Clayton! ”’ it affirms that the writer has examined a
specimen collected by the person to whose name it is appended.

— Between two figures, as in “Stamens 5-10,” indicates the extremes of
difference, as that the stamens are from five to ten.

°//’ The signs for degrees, minutes, and seconds, as 1°, 2’, 3”, are used in

Gray’s Manual of Botany of the Northern United States, for feet (°),
inches (’), and lines (’’). With European authors, usually the sign
for minutes is for feet; that of seconds for inches: thus 1’, a foot
high ; 1”, an inch long; and 1”, a line long.

O= Cotyledons accumbent to the radicle.

©]! Cotyledons incumbent on the radicle.

~ Bw BPD D UMULtEFR

GLOSSARY,

OR

DICTIONARY OF BOTANICAL TERMS, ENGLISH
AND LATIN,

COMBINED WITH AN INDEX.

Tuts Glossary is intended to contain all the principal technical terms (substan-
tive as well as adjective) of structural and systematic Botany, as far at least as
concerns Phzenogamous 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 umnecessary 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 | Abruptly pinnate. Pinnate without a

words of Greek derivation, signifies terminal leaflet or appendage; 101.
the absence of the organ mentioned; | <Acanthdcladous (-us). Having spiny
as, Apetalous, destitute of petals; branches.

Aphyllous, leafless. In words be- | Acanthéphorous (-us). Spine-bearing.
ginning with a vowel, this prefix is | Acaulescent (-ens). Stemless, or appar-

changed to an; as, Ananthous, flow- ently so, with no proper caulis; 45.

erless ; Anantherous, antherless. Acaulis. Stemless; same as Acaulescent.
Abbreviations, 385. Accessory. Something additionai, or of
Aberrant. Wandering, applied to spe- the nature of appendage.

cies, genera, &c., which differ in some Accessory Buds, 44.

respect from the usual or normal char- | Accessory Fruits, 300.

acter of the group they belong to. Accrescent (-ens). Increasing in size
Abnormal (Abnormis). Differing from with age, as often occurs with the

the normal or usual structure. calyx after flowering.
Aboriginal. Strictly native; indigenous. Accrete (-us). Grown together, or con-
Abortion (Abortus). Imperfect develop- solidated with some contiguous body.

ment or non-development of anorgan; | Accumbent (-ens). Lying against an-
179, 187. | other body.
Abortive (-ivus). Defective or barren. Accumbent Cotyledons. With edges
Abrupt (Abruptus). Terminating sud- | against the radicle: 313.
denly ; the opposite of tapering. Acephalous (-us). Headless.

5904

Acerose (-dsus). Needle-shaped, like the
leaves of Pines.

Acetabuliform (-ormis). In the form of
a shallow open cup or saucer.

Achen.um or Achenium. A small, dry
and hard, one-celled, one-seeded, inde-
hiscent fruit; strictly one of a single
and free carpel; but extended tu simi-
lar ones of more than one carpel, and

also with adnate calyx; 294. (Ache-
nium is etymologically the proper
orthography; but achenium is De-

coming the commoner form.)

Achenocarp (-arpium). General name
of a dry and indehiscent fruit ; 292.

Achenodium. Such a double achenium
as that of Umbellifere; a Cremo-
carp.

Achlamydeous (-eus).
anth; 191.

Acicula. <A bristle.

Acicular (-aris). Bristle-shaped, or slen-
der needle-shaped.

Acinaciform (-ormis). Seymitar-shaped ;
curved with rounded point, thicker on
the straighter edge than on the con-
vex edge.

Acinosus. Like grapes or grape-seed.

Acinus. Classically a berry, particu-
larly a grape, or its stony seed, or a
bunch of berries; now sometimes ap-
plied to the separate carpels of an
aggregate baccate fruit, or to the con-
tained stone or seed; 297.

Acorn. Fruit of the Oak.

Acotylédon, p|. Acotyledons, Acotylédones.
A plant or plants destitute of coty-
ledon, or

Acotyledonous (-eus). Without cotyle-
dons; as the embryo of Cusecuta; 26,
38. Mostly applied, as by Jussieu.
to plants which have no proper seed
nor embryo, and therefore no cotyle-
don; 339.

Acramphibry1. Plants producing side as
well as terminal buds or growths ; 341

Acrobrya. Plants growing from apex

only; 341.

-Acrogen (Acrdégene). Name of class of
plants which in growth are said to be

Acroyenous. Growing from the apex
or by terminal buds only.

Acrosércum. Desyaux’s name for a
berry from an ovary with adnate
calyx.

Acrospira. An old name of the plu-
mule of a grain in germination.

Aculeate (-eatus). Prickly; beset with
aculei.

Destitute of peri-

GLOSSARY.

Aculeosus. Abounding with prickles.

Aculeolate (-atus). Beset with diminu-
tive prickles, or

Aculeoli. Diminutive of aculei.

Aculeus. A prickle; a pointed small
excrescence of the bark.

Acumen. <A tapering point.

Acuminate (-atus). Ending in a tapering
point; 96.

Acute (Acutus). Terminating in an acute
angle; 97.

Acropetal. Developing from below up-
ward, or from base toward apex.

Actinomorphous (-us). Capable of bi-
section through two or more planes
into similar halves, as is a regular
symmetrical flower; 175.

Acutiisculus. Somewhat acute; acutish.

Adelphous (-us, Adelphi, brothers). Sta-
mens with cealescent or clustered fila-
ments are monadelphous, diadelphous,
&c., according to the number of Adel-
phia or brotherhoods.

Aden. Greek fur gland, is compounded
with Greek words with this meaning;
as, Adendphorus, gland-bearing; -Ad-
enophyllus, leaves bearing glands, &c.

Adylutinate (-atus). Same as accrete.

Adherent (Adherens). Generally same
as adnate; may refer to adhesions not
congenital.

Adnate (-atus). Congenitally united to ;
as the calyx-tube of the gooseberry to
the ovary; 182. Adnate anther is one
seemingly borne on the outer or inner
face of the filament ; 7. e. extrorsely or
introrsely fixed by its whole length to
the connective; 253.

Adnation. The state of being adnate;
179; 181.

Adpressus. Latin of appressed.

Adscendens. Latin of ascending.

Adsurgens. Latin of assurgent.

Adventitious, Adventive. That which has -
come from abroad or as astranger; as
a plant lately or by chance introduced
from another country.

Adventitious Buds; 45.

Ayquilateralis. Equilateral, equal-sided.

Aqualiflérus. When all the flowers of
the same head or ciuster are alike in
form as well as character.

Ayudlis, Aquans. Equal; equalling.

Aérial roots, &e. ; 33.

Aérophytes. Air-plants; 35.
Eruginosus. Verdiaris-colored.

stival (-dlis). Relating to summer.
Estivation (-io). The disposition of the
parts of a flower in the bud; 132.

GLOSSARY.

ZEterio. A form of aggregate fruit; 300.

Athevgamia, Ethogamous, 340.

Ajinity. True and near relationship;
327, 330.

Ayamous or Agamic. Destitute of sexes.

aciiomenate Heaped or crowded
pis ee eur) jin a dense cluster,
La “but not cohering.

Aggregate Fruits. Those formed of
aggregate carpels of the same flower;
248, 301.

Agrestis. Growing in fields.

Air-plunts. Plants unconnected with the
ground; 35. >

Akene, Akenium. See Achzenium.

Ala (pl. ale). A wing. Also the side
petals of a papilionaceous corolla;
185. Has also been used in the sense
of axilla.

Alabastrum. <A flower-bud; 40.

Alar (Alaris). From ala in the sense of
axilla, therefore wxillary or in the forks.

Alate (-atus). Winged.

Albescens, Albicans. Whitened, whitish,
or hoary.

Albumen of the seed. Any deposit of
Nutritive material within the seed-
ccats, and not in the embryo; 14, 309.

Albuminous or Albuminose ( Albuminvosus).
Said of seeds provided with albumen;
13, 309.

Albirnum. Sapwood; the newer wood
of an exogenous stem; 80.

Albus. White.

Allagostémonous. With stamens alterna-
tively inserted on the torus and on the

petals.

Alliaceous (-eus). Having the smell of
garlic.

Alliance. Synonym of Cohort; 326.

Allégamy. Fecundation of the ovules of
a flower by other than its own pollen ;
cross fertilization, 216.

Alp:strine (Alpestris). Growing on
mountains below an alpine region
or one unwooded from cold.

Alpine (-inus). Growing on the higher
parts of the Alps, or (by extension of
meaning) on other mountains above
the limits of trees.

Alternate (Altérnus). One after an-
other; as of leaves placed singly
instead of in pairs (opposite) or in
whorls. Also, standing before inter-
vals; as stamens alternate with petals
instead of before them; 6, 119.

Alternative (-ivus). In eestivation, with
an inner whorl alternating with an
outer one; 134, 136.

395

Alvéolate(-atus). Honeycombed ; having
deep angular cavities (Alvéoli) sepa-
rated by thin partitions, as the recep-
tacle of cotton-thistle.

Ambitus. The ray or circumference of
a head, &c.

Ament {Amentum). A catkin, or pe-
culiar scaly spike; 150.

Amentuceous (-cus). Bearing catkins, or
catkin-like.

Amorphous (-us).
definite form.

Amphanthium. One of the (needless)
names coined for a dilated receptacle
of inflorescence.

Amphibrya. Equivalent to Monocotyle-
dones; 341.

Amphicarpous (-us).
kinds of fruit.

Amplhigamous Cryptogams, 340.

Amphigastria. Peculiar leaves (of He-

_ paticze) imitating stipules.

Amphisarca. A hard-rinded berry, or
fruit succulent within and woody or
crustaceous without, as a calibash.

Amphispermium. Link’s name for a
one-seeded pericarp which is con-
formed to the seed; an akene.

Amplitropous (-us), wrongly Amphi-
tropal. Turned both ways; applied
to an ovule with hilum intermediate
between micropyle and chalaza; 279.

Amphora. A pitcher; and the lower
part of a pyxis.

Amplectens, Amplexans, Amplexus. Em-
bracing, clasping.

Amplezicaul (-aulis). Clasping a stem,
as does the base of certain leaves.

Ampliate (-atus). Enlarged or dilated.

Ampulla. A bladder or flask-shaped
organ, as of Utricularia.

Ampullaceous (-us), or Ampulleform.
In the form of a bladder or short flask.

Amylaceous (-eus). Resembling or com-
posed of starch, or Amylum.

Amyloid. Analogous to starch.

Analogy (Analogia). Likeness in cer-
tain respects. As distinguished from
affinity, it means resemblance in cer-
tain respects on/y, not in the plan of
structure. Thus, a Ranunculus is
analogous to a Potentilla, but there
is no near affinity or relatio ship be-
tween the two. And the tendril of a
Pea, that of a Smilax, and that of
the Grape-vine are analogues ; t. € , are
analogous organs, but are not homo-
logues ; for the first answers to a leaf,
the second to stipules, and the third

Shapeless; of in-

Producing two

396

toastem. The spur of a Larkspur is
analogous to one of the five spurs of
Columbine, but not homologous with
it; for the first is a sepal, and the
second a petal.

Anandrous. Destitute of stamens.

Anantherous (-us). Destitute of anthers.

Aninthous (-us). Flowerless.

Anastomosis. The connection of veins,
&c., by eress-veins, forming reticu-
lation.

Andtropous (-us), wrongly Anatropal.
The reversed ovule, with micropyle
close by the side of the hilum, and
chalaza at the opposite end; 279.

Anceps, Engl. <Ancipital. Botanically
always used in the sense of two-
edged.

Ander, andra, andrum.
pounds, the male.
Andro-diecious. With flowers on one
plant hermaphrodite, and on another

staminate only; 191.

Andrecium. The stamens of a flower
collectively ; 165, 249.

Androyynous (-us). Said of an inflores-
cence composed of both male and
female flowers.

Androphore (Andréphorum). A sup-
port or column on which stamens are
raised.

Androus. See Ander.

Aneméphilous. Literally wind-loving.
Said of flowers which are fecundated
by wind-borne pollen; 217.

Anfractuosus. Abruptly bent hither and
thither, as the stamens of Cucur-

" bita.

Angiocarpous (-us). When a fiuit is
covered by some envelope.

In Greek com-

Angiospermia. A Linnzan artificial
order; 337.

Angiospermous, Angiosperme, Anyio-
sperms. Plants with seeds borne ina

pericarp; 259.
Angular Divergence of leaves; 123.
Anisémerous (-us). Unequal in number
in the different circles of the flower;
unsymunetrical.
Anisopélalous (-us). With unequal petals.
Anisuphyllous (-us). | Unequal-leaved;
i. €., the two leaves of a pair unequal
Anisostémonous (-us) When the sta-
mens are not of the number of the
petals.
Annotinus.
growths.
Annual (Annuus)
duration; 30.

A year old, or in yearly

Of only one year’s

GLOSSARY.

Annular (-aris). In the form of a ring;
or marked transversely by rings.
The latter more properly.

Annultte (-atus). Marked with rings.

Annulus. A ring, such as that with
which the sporangia of some Ferns
aud Mosses are furnished.

Anophytes (Anophyta) Name of group
comprising Mosses, &e.

Antepositio: Same as Superposition;
179, 195.

Anterior, as to position, denotes the front
side, or averse from the axis of inflo-
rescence; 160.

Anthela. A deliquescent and paniculate
cyme, with median ramification, and
the lateral axes overtopping the
central, as in Juncus tenuis, &e.
May be either a Drepanium or a Rhi-
pidium.

Authemy, Anthemia.
of any kind; 144.

Anther (Anthera). The polliniferous
part of a stamen; 165, 251. :

Antheridium. An analogue of the an-
ther in Cryptogams.

Antheriferous (-us). Anther-bearing.

Anthesis. The time at which a flower
is perfected and opens; or the act of
expansion of a flower.

Anthocarpous(-us), Anthocarpuum. Fruits
in which some organ exterior to the
pericarp is concerned ; 300.

Anthoclinium. Name of a receptacle of
inflorescence, such as that of Com-
posite.

Anthodium. A name for the head of
flowers (or so-called compound flowers)
of Composite; 147.

Antholysis. A retrograde metamor-
phosis of a flower, in which normally
combined parts are separated.

Anthophore (Anthéphorum). The stipe
when developed between calyx and
corolla; 212.

Anthus or Anthos.
compounds.

Anticous (Anticus).
253.

Antitropous (-us), less properly Antitro-
pal. Said of an embryo with radicle
pointing to the end of the seed oppo-
site the hilum; 312.

Antrorse. Directed upward or forward.

Apetalous (-us). Having no petals; 190.

Apex. Besides its ordinary meaning,
the top of a thing, it was once the
technical name of an anther; 166.

Aphyllous (-us). Leafless.

A flower-cluster

A flower, in Greek

Facing anteriorly ;

GLOSSARY.

Apical (-alis). Relating to the apex or tip.

pices. The name for anthers anterior
to Ludwig and Linnzus; 166.

Apiculate (-us). Ending in a short
pointed tip or apicula.

Apocarpous (-us). When carpels of a
gyucecium are separate; 261, 262.

Apéphysis. An enlargement or swelling
of the surface of an organ at some par-
ticular part.

Apothecia. ‘The ‘shields ”’ or fructify-
ing disks of Lichenes.

Apotropous (-us). Said of an anatropous
ovule which when pendulous has
rhaphe averse ; 282.

Appendage, Appendix. Any superadded
or subsidiary part.

Appendiculate (-atus). Furnished with a
small appendage (Appendiculum), or
with any appendage.

Appositus. Placed side by side.

Appressed (Lat. Adpressus). Lying flat
against or together for the whole

length.
, . . .
Apricus. Growing in dry sunny places.

Apterous (-us). Wingless; not alate.
Aquatic (-icus). Living in water.
Aquatilis. Living under water.
Arachnoid (-oideus). Cobwebby ; com-
posed of slender entangled hairs.

Araneose (-osus), Araneus. Like spider-
web; same as Arachnoid.

Arbor. <A tree; 50.

Arboreous ( Arboreus).
lating to a tree.

Arborescent (-ens). Tree-like; approach-
ing the size of a tree.

Arboiétum, also Arbustum. <A_ place
where trees are grown; an arranged
collection of trees.

Arbiscula. A small shrub of tree-like
growth or form.

Arbuscularis. Ramified like a little tree.

Archegonium. The spore-case of mosses,

_&c., in an early state.

Arcuate (-atus). Moderately curved, as
if bent like a bow.

Areéola, pl. Areole. Spaces marked out
on a surface, as by the reticulation of
veins, &e.

Aréolate (-atus). Marked with areole.

Arenosus, Arenarius. Growing in sand
or sandy places.

Argentate (-atus, Argénteus). Silvery,
or shining white with a tinge of gray.

Argillosus. Growing in clayey soil.

Argos. Greek for pure white, used in
compounds; as, aryophyllus, white-
leaved.

Tree-like, or re-

397

Argitus. Sharp-toothed; said of the
serration of leaves.

Argyros. Greek for silvery; used in
compounds; as, argyrophyllus, silvery-
leaved.

Arhizal ( Arhizus). Rootless.

Avillute (-atus). Having an arillus.

Avil, Arillus. Aw extraneous or late-
formed seed-coat or covering, or an
appendage growing from or about the
hiluin of a seed; 308.

Arillifurm (-ormis). In the form of an.
arillus.

Arillode, Arillodium. <A false arillus,
or one which does not originate from
or below the hilum, but from the
micropyle or rhaphe; 309.

Arista. An awn.

Avistate (-atus). Awned; bearing an
arista.

Aristulate (-atus). Bearing a diminu-
tive awn.

Arrect (Arrectus).
upright position.

Arrow-shaped, Arrow-headed. Same as
Sagittate; 96.

Articulated (-utus). Jointed, or having
the appearance of a joint or articula-
tion ( Articulus). As of the word joint
itself, the context must show whether
the articulations mean the portions
which are connected by a joint, or the
place of connection.

Artyicial Classification, 331.

Ascending ( Adscendens). Rising upward.
Sometimes used for directed upward,
as when the stem is termed the As-
cending Axis (11); more commonly
denotes curving or rising obliquely
upward; 53.

Ascidium. A pitcher-shaped or flask-
shaped organ or appendage; 111.

Ascus. A sac; a kind of spore-cases, as
in certain Fungi and Lichenes.

Asparagi. A name for Turiones, or any
scaly shoots from underground, as
those of Asparagus.

Aspergilliform (-ormis). Brush-shaped,
i.e. like the aspergillum, or brush
used to sprinkle holy water; made up
of numerous spreading hairs, &c., in
a tuft. as the stigmas of Grasses.

Asperous (Asper). Rough to the touch.

Assimilation. The action or process by
which extraneous matter or crude food
is converted into vegetable matter.

Assurgent (Adsurgens). Rising or cury-
ing upward; 53.

Astichous (-us). Not in rows.

Brought into an

398

<Astomous (-us). Without a stoma or
mouth.

Atavism (-mus). Ancestral resemblance.

Ater. Pure black.

Athera. Greek for Arista or Awn.

Atratus. Blackened or turning black.

Atrepous (-us), wrongly Atropal. Not
turned; applied to an ovule the same
as orthotropous ; 277.

Attenuate (-atus). Slenderly tapering
or narrow.

Auctus. Same as accrescent; enlarged
after flowering ; augmented by an ad-
dition.

Augmentation. Increase beyond the
normal number; 179, 200.

Aurantiacus. Orange-colored.

Auratus, Aureus. Golden-colored, or
yellow with golden lustre.

Auricle (Auricula). An ear or ear-
shaped appendage.

Auriculate (-atus). Furnished with an
auricle; 96.

Autocarpous. A fruit consisting of peri-
carp alone, having no adnate parts.
Autogomy. Close-fertilization, the fe-
cundation of a flower by its own pol-

len; 215, 216.

Avenius. Veinless.

Auwl-shaped. Narrow, terete or some-
what so, and attenuate from a broader
base to a slender or rigid point.

Awn. A bristle-shaped appendage, such
as the beard of Rye and Barley.

Awned. Furnished with an awn.

Azil (Azilla). The angle formed on the
upper side of the attachment of a leaf
‘with the stem, or the point just above
this attachment; 6.

Axillary (-dris). In or relating to an
axil; 7.

Azile, Azial ( Azilis).
longing to the axis.

Azis. The stem; the central part or
longitudinal support on which organs
or parts are arranged ; the central line
of any body.

Relating or be-

Bacca. A berry; 299.

Baccate (-atus). Berry-like; pulpy
throughout.

Baccetum. An aggregation of berries

in one flower; 300.

Badius. Chestnut-brown.

Balausta. Name applied to the fruit
of the Pomegranate, with firm rind,
crowned with the lobes of an adnate
calyx, baccate within, and many-
seeded.

GLOSSARY.

Banner. The vexillum, standard, or
upper petal of a papilionaceous co-
rolla; 184.

Barb. A bristle or stout hair, which is
hooked or double-hooked, or retrorsely
appendaged at the tip.

Barba. Beard.

Barbate (-atus). Bearded; beset with
long and weak hairs.

Barbellate (-atus). Beset with shorter
and stiffer hairs or barbelle.

Barbellulate (-atus). Diminutive of the
preceding. :

Bark. The rind or cortical portion of a
stem, especially of an exogen; 76.

Basal ( Basilaris). Relating to the base.

Basé@!-nerved. With nerves all from the
base of the leaf; 92.

Base (Basis). The extremity by which
an organ is attached to its support.
Basidia. Cells of the fructification of
Mushrooms which bear the spores.
Buasificed (-us). Attached by the base

or lower end; 253.

Basigynium. Synonym of Carpophore
or Thecaphore.

Basinerved (-ius). When the ribs pro-
ceed from the base of a leaf.

Busipetal. Developing from apex to-
ward the base.

Bast, or Bass. Inner fibrous bark; 77.

Bast-cells. The es-ential components of
bast; long and flexible butthick-walled
attenuated cells; 77.

Beak. A narrowed or prolonged tip.

Beaked. Ending in a beak.

Bell-shaped. Same as Campanulate ; 249.

Berried. Baccate.

Berry. A fruit, the whole pericarp of
which is fleshy or pulpy; 299.

Bi- or Bis. As a prefix to Latin words
(Greek words have Di-), two, twice,
or doubly.

Biacuminate (-atus). _Two-pointed, as
malpighiaceous hairs, fixed by the
middle and tapering to each end.

Biarticulate (-atus). Two-jointed.

Biauriculate (-atus). Two-auricled.

Bibracteate ‘-atus). With two bracts.

Bibracteolate (-atus). With two bract-
lets.

Bicallose (-osus).

Bicarinate (-atus).

With two callosities.
Two-keeled.

Biceps. With two supports or stalks,
Bicipital. ) or two-headed.
Bicolor. Two colored.

Biconjugate (-atus). Twice paired.
Bicornis. Two-horned.
Bicurnute. Saie as preceding.

GLOSSARY.

Bicruris. Two-legged, or with two sup- | ABipinnate (-atus).

ports.

Bidentate (-atus). Having two teeth.
(Not doubly dentate.)

Biluus. Lasting two days only.

Biennial ( Biennis). Of two years’ dura-
tion; 31.

Bifurious (-ius).
vertical rows.

Biferus. Double-bearing; fruiting twice
a year.

' Bifid (-idus). Two-cleft, to the middle
or thereabout.

Bifldrous (-us). Fwo-flowered.

Bifoliate. Two-leaved.

Bifoliolate. Of two leaflets.

Biferate (-atus). Having two open-
ings.

Biformis. Two-formed; in two shapes.

Bifvrons. With two faces or aspects.

Bifireate (-atus). Two-forked; i.e. of
two prongs or forks. But it may
mean bis furcatus; i.e., forked and
again forked.

Bigeminate (-atus).
as Biconjugate.

Biyener. The offspring of a cross be-
tween two generically different plants.

Bijugate (Bijugus). Two-paired, as a
pinnate leaf of two juga or pairs of
leaflets.

Bilabiate (-atus). Two-lipped; 247.

Bilamellate (-atus), or Bilamellar. Of
two plates or lamellae.

Bilobed ( Bilobus), or Bildbate. Of two

_ lobes, or cleft into two segments.

Bilocellate. Divided into two locelli;
263.

Bilocular (-arvis). Two-celled.

Bimestris. Lasting two months.

Bimus. Lasting two years; two years
old.

Binary (-arius).
members; 176.

Binate (-atus). In pairs or twos.

Bini. Twin, or two together.

Binodal ( Binddis). Having two nodes.

Binomial Nomenclature, 346.

Bivlogy. “The natural history of plants
and animals, 7. e. of living things; 1.

Bipalmate (-atus). Twice palmately com-
pound.

Biparous. Bearing two; as a cyme of
two rays or axes; 152, 155.

Bipartible (-ibilis). Capable of division
into two similar parts.

Bipartite (-itus). Divided almost into
two pieces; two-parted.

Bipes. Same as Bicruris.

Two-ranked; in two

Twice twin; same

Consisting of two

399

Doubly or twice
pinnate; 103.

Bipinnatifid (-idus).
pinnatifid; 100.

Bipinnatisect (-us). Twice pinnately di-
vided.

Biplicate (-atus). Twice folded or plaited.

Biporose (-osus). Opening by two pores.

Biradiate (-atus). Of two rays.

Birimose (-osus). Opening by two slits.

Bisected (-us). Comyletely divided into
two parts; 99.

Biseptate (-atus). With two partitions.

Biserial (-ialis), or Biseriate (-iatus). In
two series, one above the other.

Biserrate (-atus). When serratures are
again serrate: doubly serrate.

Bisexual. Having both stamens and
pistil; hermaphrodite; 191.

Bisilcute (-atus, Bisulcus). Two-grooved;
having two furrows.

Bitérnate (-atus). Twice ternate.

Bladdery. Thin and inflated.

Blade. The lamina, limb, or expanded
portion of a leaf, &e.; 85, 245.

Blastéma. The budding or sprouting
part or point. First used for the axis
of an embryo; now used for the ini-
tial growth out of which any organ
or part of an organ is developed.

Bloom. Besides its use as equivalent to
blossom, it denotes the white powdery
and glaucous covering of the surface
of many fruits and leaves, of a waxy
nature.

Boat-shaped. Of the shape of a boat, of
the deeper sort, with or without a keel.

Bostrychoidal. Having the form or char-
acter of a ringlet, or Bostryx: 157.

Twice. or doubly

Bostryz. An uniparous helicoid cvme;
156.
Bothrénchyma. Tissue of plants com-

posed of dotted or pitted ducts.
Botry-cymose. Racemes or any botrvose
clusters cymosely aggregated ; 159.
Botryose (-osus), Botryoidal. Of the ra-
cemose type; 144. 145, 146, 153.
Botrys. The equivalent of Raceme ; 146.
Botuliformis. Sausage-shaped.
Biachiate (-idtus). With spreading
arms, as branches (especially opposite
and decussate) widely diverging.
Brachys. Greek for short, and used in
compounds; as, Brachypodus, short-
stalked.
Bract, Bractea. The leaves (more or
less modified) of a flower-cluster; 118,
141.

Bracteate (-eatus). Having bracts.

400

Bractéola, Bracteole. See Bractlet.

Bracteolate(-atus). Having bractlets.

Bractlet. A bract of the ultimate grade,
as one inserted ona pedicei or ultimate
flower-stalk, instead of subtending it;
141, 142, 160.

Bracteose (-osus).
spicuous bracts.

Branches. Secondary axes, or divi-
sions of an axis; 47.

Branchlets. Ultimate branches or divi-
sions of an axis; 47.

Breathing-pores. See Stomata, 89.

Bristle. A stiff hair, or any slender
body or outgrowth which may be
likened to a hog’s bristle.

Bristly. Beset with bristles.

Brunneus. Deep brown.

Brush-shaped. See Aspergilliform.

Bryology. The botany of Mosses.

Bud. The undeveloped state of a stem
or branch, with or without leaves;
6, 40.

Bud-scales. The teguments of a bud; 40.

Bulb ( Bulbus). A \eaf-bud (commonly
subterranean) with fleshy scales
eats; 43, 62.

Bivbiceps. A stem with bulbous base.

Bulbiferous (-us). Bulb-bearing.

Bulbiilus, Builbulus. Diminutive bulb.
Same as

Bulblet. A small bulb, especially such as
is produced in the air, in the axil of or-
dinary leaves, or upon them; 63.

Bulbodium. A synonym of Corm, the
“solid bulb.”

Bulbo-tuber. Synonym of Corm.

Bulbous, Bulbosus. Having bulbs or the
structure of a bulb.

Bullate (-atus). Said of a puckered sur-
face (as if blistered), thrown into por-
tions which are convex and projecting
on one side and concave on the other.
Also used in specific names, in its
more literal sense for inflated.

Bursicula. A small pouch (bursa);
such as that which encloses the disk
or gland of the caudicle of the pollin-
ium of an Orchis.

Bursiculatus. Furnished with a bursi-
cula or pouch.

Byssaceous (-eus). Composed of fine
threads, like dyssus or fine flax.

Full of, or with con-

Cadicous (-us). Dropping off very early,
as the calyx of a Poppy at the time
of expansion; 243.

Ceruleus. Sky blue. or pure blue.

or

}

GLOSSARY.

Cesius. Lavender-color; pale green with
whitish or gray.

Calathidium, Calathis. Literally a bas-
ket; aname for the head of flowers (or
better for the involucre only) of Com-
positee

Calathiform (-ormis). Cup-shaped ; of
somewhat hemispherical outline.

Calcar. A spur; mostly used for the nec-
tariferous one of a calyx or corolla.

Calcarate (-dtus). Furnished or pro-
duced into a spur.

Calceolate (-atus), or Calceiformis.
Shaped like a slipper or shoe.

Callose (-osus). Bearing callosities
(calli), or hard protuberances.

Calvus. Bald, as an akene without
pappus.

Calycanthemy. Name of the monstros-
ity in which the calyx imitates an
exterior corolla: 174.

Calyciflorous (Calyciflore), 340.

Calycine (Calycinus). Relating to calyx.

Jalyculate (-atus). Bearing bracts next
the calyx which imitate an external
or accessory calyx.

Calyculus. An involucre or involucel
imitating an additional calyx.

Calyptra. The hood or veil of the
spore-case of a Moss; or some cover-
ing body like it.

Calyptrate (-atus). Furnished with a
calyptra, or something like it.

Calyptriform (-ormis). Calyptra-shaped ;
as the calyx of Eschscholtzia.

Calyz. The flower-cup, the exterior
perianth ; 164.

Camara and its diminutive Camérula
(chamber) are sometimes used for the
cells of a fruit.

Cambium. Old name of the viscid mat-
ter between bark and wood in com-
mon trees or shrubs in spring; now
used for the nascent structure there
forming, or Cambium layer ; 78.

Campanulate (-atus). Bell-shaped ; elon-
gated cup-shaped or shorter, and broad
from the base; 249.

Campaniformis. Same as Campanulate.

Campylosp:rmous (-us). Curved-seeded.
Said of seed-like fruits or carpels, as
those of some Umbellifere, in which
the contained seed is involute by the
lateral edges, so as to produce a longi-
tudinal furrow on the ventral face.

Campylotropous (-us), or less correctly
Campylotropal, or Campulitropous.
An ovule or seed which is curved
in its formation so as to bring the

GLOSSARY.

micropyle or true apex down near to
the hilum; 279.
Canalicaulate (-atus). Channelled, or
with a longitudinal groove.
Cancellate (-atus). Latticed; resembling
lattice-work.

Candidus. Pure white.

Canescens. Hoary, usually with gray
pubescence.

Canus. Gray-white; whiter than the

preceding.

So slender that it

may be compared

with the hairs of
animals.

Capitate (-atus). WHead-shaped, or col-
lected in a head; 147.

Capitellate (-atus). Diminutive of Cap-
itate.

Capitulum. A head of or simple globu-
lar cluster of sessile flowers; 147.

Capreolate (-atus). Bearing a tendril
(capreolus).

Capsule (-ula). A dry and dehiscent
pericarp composed of more than one
carpel ; 289, 293.

Capsular. Of the nature of, or relating
to, a capsule.

Capsuliferous. Capsule-bearing.

Carcvulus. An unused name for an
indehiscent and several-celled dry
fruit; 297.

Carina. <A keel; used either for the
two combined lower petals of a papil-
ionaceous corolla (185); or for a sa-
lient longitudinal projection on the
centre of the lower face of an organ,
as on the glumes of many Grasses.

Carinate (-alus). Keeled.

Cariopsis or Caryopsis. A grain; a seed-
like fruit with thin pericarp adnate to
the contained seed ; 295.

Carneus. Ylesh-colored, very pale red.

Caro. Flesh, as the pulp of a melon, or
the fleshy part of a drupe.

Carpadilium. Synonym of Cremocarp.

Carpel, Carp/llum. A simple pistil, or
an element of a compound pistil, an-
swering to one leaf; 167, 260.

Carpid, Carpidium. Synonym of carpel.

Carpology. The botany of fruits.

Carpophore (Carpéphorum). A portion
of receptacle prolonged between the
carpels; 212.

Carpophyll (Carpophyllum). Literally
fruit-leaf; svnonym of Carpel; 260.
Cartilaginous or Cartilagineous (-eus).
Of the texture of cartilage or gristle ;

firm and tough.

Capillaceous (-eus).
Capillary (-aris). F

401

Caruncle (Carincula). An excrescence
at or about the hilum of certain seeds ;
308.

Caryophyllaceous(-eus). Resembling or
relating to the corolla of Dianthus
Caryophyllus (246), or to the Pink
family.

Caryopsis. See Cariopsis.

Cassideus. Helmet-shaped.

Cassus. Empty, as an anther contain-
ing no pollen.

Castrate (-atus). Said of a stamen
which wants the anther.

Catapetalous (-us). Where petals are
united only by cohesion with united
stamens, as in Mallow.

Catuphylla. Answers to the German
‘* Niederblatter,’’ or under-leaves,
those at the beginning of a growth,
cotyledons, bud-seales, scales on rhi-
zomes, &c.; 6.

Catenulate (-dtus). Formed of parts
united end to end, like the links of a
chain.

Catkin. A scaly spike (see Ament); 150.

Caudate (-atus). Furnished with a tail
(cauda), or with a slender tip or ap-
pendage resembling a tail.

Caudex. A trunk or stock of a plant: 50.
Caudicle (Caudicula). The stalk of a
pollinium, &e.
Caulescent (-ens).

stem.

Cailicle (Cauliculus). The initial stem
in an embryo, generally named the
Radicle; 10.

Cauline (-inus). Belonging to the stem.

Caulis. Greek form Caulon. The stem
of a plant.

Caulocaipic or Caulocarpous. Applied
to plants which live to flower and
fructify more than once or indefi-
nitely.

Cauléme, Cauloma.
plant.

Cephalanthium. One of the names of
the head or capitulum in Composite ;
148.

Cell ( Cellula).

Having an obvious

The stem-part of a

The anatomical element
of plants; 28. The cavity of an
anther which contains the pollen,
or an anther-lobe, thus taken in the
sense of the circumscribing wall as
well as the cavity; 251, 254. The
cavity, or any one cavity of an ovary
or pericarp, containing the ovules or
seeds; 262.

Cellular Plants, Cellulares, 340.

Céllule (-ula). Diminutive of cell; of

402

the same meaning as Cell in vegetable
anatomy; 28.

Cellulose. The material, chemically con-
sidered, of which the wail of the cell
consists.

Cenchium. A name of the peculiar four-
parted fruit (or the four nutlets around
a common style) which distinguishes
Labiate and Borraginacee.

Centiifjugal. Tending or developing
from the centre outward.

Centripetal. Tending or developing from
without toward the centre.

Cephalanthium. Synonym of Antho-
dium.

Ceratium. A siliquiform capsule, such
as that of Corydalis, Cleome, &c.

Cereal. Belonging to corn and the
allied grains.

Cerinus. Of the color of wax.

Cérnuous (-uus). Nodding.

Chet. Greek for a bristle, Latin Seta.

Chaff. Small scales; dry and depau-
perate bracts; such as those on the re-
ceptacle of asunflower and many other
Compositze ; also glumes of Grasses.

Chaffy. Provided with or having the
texture of chaff.

Chalaza. The part of an ovule where
coats and nucleus are confluent; 277.
Channelled. Hollowed out longitudi-

nally like a gutter. See Canaliculate.

Character. A diagnostic description,
or the enumeration of essential differ-
ences; 361.

Chasmoyamy. The opening of the peri-
anth at flowering time; the opposite
of Cleistogamy

Chartaceous (-eus)
of writing-paper.

Chlorophyll. The green matter of leaves
and other vegetation ; 76, 88.

Chloros. Greek for green. Enters into
compounds, such as Chlorunthus,
green-flowered, Chloranthy, same as
Chlorosis, as when petals turn green;
12.

Chlordsis. Literally becoming green, as
some flowers in retrograde metamor-
phosis. Also used contrariwise for
the loss of a normal green color; 172.

Chorda Pistillaris. A line of tissue reach-
ing from stigma to ovary.

Choripetalous (-us). Same as Polypeta-
lous, 7. e. petals unconnected : 244.

Chorisepalous. Same as Polysepalous.&e.

Chorisis. The separation of a leaf or
phyllum into more than one ; 202.

Cher’stophyllus. Separate-leaved.

Having the texture

GLOSSARY.

Chromule (-ulz). Coloring matter of
plants other than chlorophyll, espe-
cially that of petals.

Chrysus. Greek for golden, or golden-
yellow; as
Chrysanthus. Yellow-flowered, &c.

Cicatrixz, Cicatricula. A scar left by
the fall of a leaf or other organ.

Ciliate (-atus), Ciliaris. Marginally
fringed with hairs.

Cilium, pl. cilia. Marginal hairs, form-
ing a fringe, like the eyelash. (The
name has been extended in scientific
books to undividual hairs, and of a
surface as well as edge.)

Cincinnus. A’curl: name of a uniparous
scorpioid cyme, which is Cineinnal ;
156, 157.

Cinenchyma. Laticiferous tissue.

Cinerascens, Cincraceus. Ash-grayish.

Cimereous (-eus). Ash-gray.

Cinnabarinus. Cinuabar-color ; searlet
touched with orange ‘

Circinal (-alis). Involute from the tip
into a coil; 133.

Circinate, or Circinnate (-atus). Same
as preceding; or sometimes meaning
coiled into a ring only.

Circumscissile, or Circumcissile ( Cireum-
scissus). Cut cireulary and transverse-
ly; divided transversely ; 231, 293.

Circumscription (-iv). The general out-
line of the margin of a flat body.

Cirrhiferous (-us) and Cirrhose.
dril-bearing.

Cirrhus. A tendril; 54.

Citreus, Citrinus. V.emon-colored.

Clados. Greek for branch; whence such
terms as

Cladodium. Same as Cladophyllum.

Cladophyll, Cladophylla. Branches as-
suming the form and function of foli-
age: 65, 66.

Ten-

Claivate (-atus), Claviformis. Club-
shaped.
Clivellate. Diminutive of Clavate.

Claviculate (-atus). Furnished with clar-
icule ; viz, tendrils, hooks, or other
appliances for climbing.

Class, 325.

Classification, 315.

Clathrate (-atus). Latticed.

Claw. The narrowed base or stalk which
some petals, &c., possess; 245.

Cleistégamy, Cleistogamous, Cleistogamic.
Close-fertilization in unopened blos-
soms ; 241.

Cleistégeny, Cleistogenous, 241. Same as
Cleistogamy.

GLOSSARY.

Cleft Cut half-way down or there-
about; ¥8.

Climbmy. Rismg by laying hold of
surruunding objects for support; 51.
Clinandrium. ‘The anther-bed in Orchi-

dacex.

Clinanthium. A name for the receptacle
of inflorescence in Composite; 148.
Clinium. Used in Greek compounds for
receptacle, e.g. Periclinium, for an
involucre around the receptacle of

inflorescence.

Close-fertilization. Fecundation by own
pollen ; 216, 280.

Cloves. A gardener’s name for young
bulbs developed around a mother bulb.

Club-shaped. Gradually thickened up-
ward from a slender base.

Clustered. Collected in a bunch of any
sort. Cluster is a good indefinite name
fur any assemblage of flowers on a
plant.

Clypeate (-atus), Clypeiformis.
ler-shaped.

Coucerrate (-atus).

Buck-

Heaped together.

Coddnate (-atus), Coadunatus. Same as
Adnate.

Coalescence. Union of similar parts;
179, 180.

Coalescent (-ens), Coalitus. Cohering;
properly applied to the organic cohe-
sion of similar parts.

Coarctate (-atus). Crowded together.

Coated. Composed of layers as an
onion, or furnished with a covering
or rind.

Cobwebly. Bearing long and soft entan-
gled hairs.

Coccineus. Bright red or scarlet (red
with a little yellow).

Coccus. Greek for a kernel or nutlet,
from which the Latin Coccum, the
kermes or scarlet grain (supposed
berry) of the Quercus coccifera; used
botanically, mostly in the form of
“coccus,”’ for the portions into which
a schizocarp, or lobed fruit with one-
seeded cells, splits up; these portions
are Cocci or Coccules; 296.

Cochlear (Cochlearis). Spoon-shaped.
Unmeaningly applied also to a form
of imbricative sxstivation with one
piece exterior; 137.

Cochleate (-atus). Shell-shaped, i. e.
spiral in the manner of a snail-shell.
Celospéermous (-us). Hollow-seeded; ap-
plied to seed-like carpels of Umbelli-
ferze with ventral face incurved at top

aud bottom, as in Coriander.

403

Cenanthium. Synonym of Clinanthium.

Cenobio. Synonyin of Carcerulus.

Cohesion. ‘Vhe congenital union of one
organ with another; either similar
parts (coalescence), or dissimilar parts
(adnation ).

Cohort. In classification a group next
superior to order, 326.

Coleorhiza. Root-sheath; the invest-
ment (belonging to the cotyledon or
plumule) through which the primary
root in many Monocotyledons bursts
in germination; 26.

Collar (Cullum). Name of an imaginary
something intermediate between pri-
mary stem and root.

Collateral. Standing side by side.

Collective Fruits. The aggregation of
the fruits of several flowers into one
mass; 301.

Colored. Of other color than the green
of herbage; 118.

Columella. The persistent axis of cer-
tain capsules, spore-cases, &c.; 289.
Column (Columna). Body formed by
the union of the filaments among them-
selves (as in a malvaceous flower), or
with the style or stigma, as in Or-

chids; 290.

Columnar. Column-shaped ; _ pillar-
shaped.
Coma. Literally a head of hair; a tuft

of hairs of any sort ; specially a tuft of
hairs on a seed: 306 Also the name
of the whole head of a tree.

Commissure (-ura). The face by which
two carpels cohere, as in Umbellifere.

Common ( Communis). General or prin-
cipal, as opposed to partial.

Coumose (-osus), sometimes
Furnished with a coma.

Cémplanate (-atus). Flattened.

Complete (Completus). Having all the
parts belong to it or to the type; 175.

Complicate (-atus). Folded upon itself.

Compound. Said of similar parts aggre-
gated into a common whole. Com-
pound Flower, 147. Compound Pistil,
263. Compound Inflorescence, 159.

Compound Leaf. One divided into sep-
arate blades; 100.

Compressed (-us). Flattened lengthwise.

Concaulescence. A name for the coales-
cence of axes: 158.

Conceptacle (-aculum). Originally used
by Linneus for what is now called
Follicle; and later for the pair of fol-
licles of Asclepiadacee and Apocy-
nacez.

Comatus.

404

Conchiformis. Shaped like one valve of
a bivalve shell.

Concinnus. Neat or elegant.

Concolor. Of the same or of uniform
color.

Conduplicate (-atus), Conduplicativus.
Volded together lengthwise; 133.

Cone. See Strobile.

Conjertus. Closely packed or crowded.

Conferruminate (-atus). Stuck together
by adjacent faces, as the cotyledons of
Horsechestuut; 314.

Confluent (-ens). Blended into one; pass-
ing by degrees the one into the other.

Conformed (-ormis). Similar to in form;
or closely fitted to, as a seed-coat to
the nucleus.

Congested (-us). Crowded together.

Conglobate (-atus). Collected into a ball.

Conglomerate (-atus). Densely clus-
tered or heaped together.

Coniferous (-us). Cone-bearing.

Conjugate (-atus). Coupled; in single
pairs. Conjuyate-pinnate, 104.

Connate (-atus). United congenitally;
107, 182.

Connate-perfoliate. United at base in
pairs around the supporting axis; 108.

Connective (-ivum). A portion of a sta-
men which connects the two cells or
lobes of an anther; 251.

Connivent (-ens). Coming into contact
or converging.

Conocarpium. An unused name for an
aggregate fruit, such as a strawberry,
consisting of many carpels on a coni-
cal receptacle; 238

Consolidated. When unlike parts are
coherent.
Continuous. The reverse of articulated

or interrupted.

Contorted (-us). Twisted; or bent or
twisted on itself. In /Estivation, the
same as Conyolute; 138.

Contortuplicate (-atus).
plaited or folded.

Contracted. Either narrowed or short-
ened.

Contrary (-arius). Opposite in direction
to the part compared with; asa silicle
compressed contrary to the dissepi-
ment.

Convolute (-utus) or Convolutive (-ivus).
Rolled up from the sides or longitudi-
nally. In stivation, 138. In Ver-
nation, 133.

Céralloid (-eus). Coral-like.

Céreulum. Old name for the embryo,
or Cor seminis; 311.

Twisted and

GLOSSARY.

Cordate (-atus), sometimes Cordiform
(-ormis). Heart-shaped; like the fig-
ure of a heart on cards; the stalk at
the broader and notched end ; 96.

Coriaceous (-eus). Leathery in consist-
ence.

Cork, 81.

Corky. Of the texture of cork.

Corky Envelope, 76.

Corm (Cormus). A bulb-like fleshy
stem, or base of a stem; a “solid
bulb;” 61.

Cormophytes ( Cormophyta), 341.

Corneous(-eus). Of the texture of horn.

Corniculate (-atus). Furnished with a
little horn.

Cornu. A horn; i.«. a horn-like process;
sometiynes used for Calcar, a spur.

Cornute (-utus). Furnished with a horn-
like process or spur.

Corolla. The interior perianth, com-
posed of petals; 165, 243.
Corollaceous (-eus), Corollinus.
ing to, or resembling corolla.
Corolliferous (-us). Bearing a corolla.

Corclliflorous, Corolliflore, 340.

Coréllula. Dimiuutive of corolla.

Corona. A crown: an inner appendage
to a petal, or to the throat of a corolla;
210, 246. Orany coronet-like append-
age at the summit of (crowning) an
organ.

Céronate (-atus).
corona, &e.

Coréniform (-ormis).
crown or coronet.

Cérrugate (-atus or -ativus).
or in folds; 133.

Cortex. Rind or bark.

Cortical (-alixs). Relating to bark.

Corticate (-atus). Coated with a bark
or with an accessory bark-like cover-
ing.

Cérymb (Corymbus). A flat-topped or
merely convex and open flower-cluster
of the indeterminate or centripetal
order; 146.

Corymbiferous (-us). Bearing corymbs.

Cérymbose. In corymbs, or in the man-
ner of a corymb. The corymb of
Linneus and of other writers down
to Roeper included most cymes. So
that much cymose inflorescence is
in descriptions loosely said to be
corymbose, or a stem is said to be
corymbosely branched, even when
the evolution is centrifugal ; 146.

Costa. _A rib; when single, a midrib
or mid-nerve. ae

Pertain-

Crowned, having a
Shaped like a

Wrinkled

GLOSSARY.

Costal-nerved. With nerves springing
from a midrib; 92.

Costate (-atus). Ribbed; furnished with
one or more lorgitudinal primary
veins or ribs.

Cotylédons ( Cotyl’don, pl. Cotyledones).
The *seed-lobes,”’ being the leaves or
first leaves of the embryo; viz., the
one, or the pair, or rarely the whorl of
leaves borne by the radicle or caulicle;
10, 311, 313. :

Cotyliformis. Dish-shaped, or wheel-
shaped with an erect or ascending
border. :

Cratéviform (-ormis). In the shape of
a goblet or cup, of hemispherical con-
tour or more shallow; 248.

Cr*mocarp (Cremocarpium). A dry and
seed-like fruit, composed of two one-
seeded carpels, invested by an epigy-
nous calyx, and separating at matu-
tity; 297.

Creeping. Running along or under
ground and rooting; 53.

Crena, Crenatura. A rounded tooth or
notch.

Crenate (-atus). Toothed by crenatures :
scalloped; 98

Crenei, Crenelled. Same as Crenature
and Crenate.

Crénulate (-atus). Diminutive of Cre-
nate, 2. ¢. with small crenatures.

Crested. Yurnished with any elevated
line, ridge, or conspicuous elevation
on the surface, especially such as may
be likened to the crest of a helmet.

Cretaceus. Chalk-white; chalky.

Cribrose (-osus) and Cribriform (-ormis).
Pierced like a sieve.

Cribriform Cells, 77.

Crinitus. Bearded with long and weak
hairs.

Crispatus. Curled or crispy.

Cristate (-atus). Crested.

Croceus, Crocatus. Safiron-colored, 7. e.
deep reddish-yellow.

Cross-breeds. The progeny of interbred
varieties; 321.

Cross-fertilization. Fecundation by pol-
len of another flower and of another
individual; 216.

Crown. See Corona, 210; 246.

Crowned. See Coronate.

Crowning ( Coronans).
suinmit of an organ.

Cruciate (-atus), Cruciform (-ormis).
Cross-shaped.

Cruciferous (-us). Cross-bearing: used
in the sense of Cruciform; as the

Borne on the

1

405

“‘cruciferous’’ corolla of the order
Crucifere ; 246.

Crumpled. See Corrugate.

Crustaceous (-us)., Of hard and brittle
texture.

Cryptos. Greek for concealed; whence

Cryptogamia. Cryptogamous or Cryp-
togamic plants ; 3, 335, 544.

Cryptégamous. Pertaining to the above.

Cucullate (-dtus), Cucullaris, Cuculli-
Jormis. Hooded, or hood-shaped,
cowled.

Culm (Culmus). The peculiar stem or
straw of Grain-plants and Grasses; 40.

Cultrate (-atus), Cultriformis. Shaped
like a broad knife-blade. -

Cuneate ( Cuneatus), Cuneiform (-ormis).
Wedge-shaped; triangular with an
acute angle downward; 95.

Cup-shaped. In the form of a drinking-
cup.

Cupule (Cupula).
the iike; 296.
Cupularis, Cupulatus. Furnished with
or subtended by a cupule or any re-

sembling body.

Cupuliferous (-us). Cupule-bearing.
Curvinerved (-ius). When the ribs of a
leaf are curved in their course; 92.
Curviserial. In curved or oblique ranks;

124.

Cushion. The enlargement at or be-
neath the insertion of many leaves.
Cuspidate (-atus). Tipped with a Cusp,

or sharp and rigid point; 97.

Cut. Same as incised, or in a general
sense as cleft.
Cuticle ( Cuticula).

or pellicle.

Cutting. A severed portion of a plant
used for bud-propagation: 43.

Cydneus. A clear bright blue.

Cyathiform (-ormis). Cup-shaped; in the
form of a

Cyathus. A drinking-cup, such as a
goblet or wine-glass.

Cycle. A circle. Sometimes used for
one turn of a helix or spire; 122.

Cyclical. Relating to acycle; or coiled
into a cirele; 119, 120.

Cylindraceous. Somewhat or nearly
cylindrical.

Cyl: ndrical (-us). Elongated and with
circular cross-section; in the form of a
cylinder.

Cymbeform or
Boat-shaped.

Cyme ( Cyma).
determinate

The acorn-cup and

The outermost skin

Cymbiform (-ormis).

A flower-cluster of the
or centrifugal type,

406

especially a broad and flattish one;
151.

Cymo-bétryose. When cymes are ar-
ranged in botryose manner; 159.

Cymose (-osus). Bearing: cymes, or re-
lating to a cyme; 181.

Cymule (Cymula). Diminutive ecyme,
or a portion of a cyme; 151.

Cynarrhodium. Name of such a fruit as
that of the Rose; fleshy, hollow, and
euclosing achenia.

Cypsela. Name of an ache: ium in-
vested by an adnate calyx, as the fruit
of Composite; 295-

Cystclith. Onz of the mineral and
usually partly crystalline concretions
of the cells of the epidermis of or
subjacent tissue of the leat in various
plants, especialiy in Urticacez.

Cytoblast. An obsolete name for the
nucleus of a cell of cellular tissue.

Dactylose (-osus). Fingered, or finger-
shaped.

Dasyphyllous (-us). Woolly-leaved.

Deald ite (-atus). Whitened over (as if
whitewashed) with a white powder or
minute pubescence.

Deca. Greek for ten, compounded with
various words, such as

Deengynia. One of the Linnean artificial
orders; 337.

Decayynous (-us).
carpels.

Decamerous(-us). Of ten members; 176.

Decandria. A Linnean class with ten
stamens: 334.

Decandrous (-us).
249.

Decapétalous (-us), Decasepalous, &c.
With ten petals or sepals, &c.

Deciduous (-us). Falling, or subject to
fall in season, as petals after anthesis,
and leaves (except of evergreens) in
autumn; 245.

Déclinate (-atus), or Declined. Bent or
curved downward or forward.

Decompound. Several times compound-
ed or divided; 102, 104.

Decompositus. Decompound.

Decumbent (-ens). Reclining, but with
summit ascending; 53.

Decurrent (-ens), Decursive. Running
down into; as where leaves are seem-
ingly prolonged below their insertion,
and so run down the stem.

Decussate (-atus). In pairs alternately
crossing at right angles.

With ten styles or

With ten stamens;

GLOSSARY.

Deduplication, Fr. Dédoublement. Same
as Chorisis; 202.

Definite (-itus). Of a fixed number, not
exceeding twenty ; or of a fixed or. er.

Definite Inflorescence. Where axes of
inflorescence end ina flower; 144, 151.

Dejflexed (-us). Bent or turned abruptly
downward.

Dejflorate (-atus).
state.

Defoliate (-atus). Having cast its leaves.

Defoliution, 87. :

Dehiscence (-entia). The mode of open-
ing of a capsule or anther by valves,
slits, or regular lines; 288.

Dehiscent (-ens). Opening by regular
dehiscence; 292.

Deliquescent (-ens). Dissolving or melt-
ing away, as a stem divided into
branches; 48.

Deltoid (-oides). Having the shape of
the Greek letter A.

Demersed (-us). Under water; same as
submersed.

Dendritic (-icus), Dendroid (-oideus).
Tree-like.

Dendron.

Past the flowering

Greek for tree.

Deni. Ten together.

Dens. A tooth.

Dentate (-atus). Toothed; specially-with
salient teeth not turned forward ; 98.

Denticulate (-atus). Minutely toothed ;
having denticulations, or diminutive
teeth.

Denidate (-atus). Made naked; stripped.

Deorsum. Downwards.

Depauperate (-atus). Tmpoverished; as
if starved; or diminutive for want of
favorable surroundings.

Depressed (-us). Having the appear-
ance or shape as if flattened from
above.

Derma. Greek for skin or surface of a
plant or organ.

Descending (-ens). Tending or turning
gradually downward.

Descending Axis. Primary root; 11.

Determinate. Limited in number or ex-
tent; as are the axes of determinate
inflorescence; 144, 151.

Désinens. Terminating in.

Desmus. Greek for things bound, or as if
chained together.

Dextrorse (Dextrorsus: adv. Deztror-
sum). Toward the right hand, or re-
lating to it; 51, 140.

Di, Dis. In Greek compounds, two, or
double.

Diachenium. Synonym of Cremoearp.

GLOSSARY.

Diadelphia. A Linnean (335) class
having the stamens.

Diadelphous (-us). Combined by their

_ filaments into two sets; 250.

Diagnosis. A brief distinguishing char-
acter.

Dialypetale, 341.

Dialypetalous (-us). Same as poly peta-
lous, t. €. of separate petals; 244.

Dialyphyllous (-us). Bearing separate
leaves.

Diandria. A Linnean class with per-
fect flowers having only two stamens;
334. si

Diandrous ( Diander, &e.). Having two
stamens; 249.

Diaphanous (-us).
shine through.

Dicarpellary. Composed of two carpels
or pistil-leaves; 261

Dichasium. A two-parted or two-rayed
eyme; 152, 155.

Dichlamydeous (-eus). Having a double
perianth; 191.
Dichétomous (-us).

two-forked.

Dichéyamous (-us), Dichogamy. Her-
maphrodite with one sex earlier de-
veloped than the other in the blossom;
219.

Dicksium. Name of a fruit consisting
of an achenium within a separate and
free covering made of perianth, as that
of Mirabilis.

Diclinvus (Diclinis). When flowers are
of separate sexes; 191.

Dicéccous (-us). Fruits of two cocci.

Dicotyl-dons, Dicotyledones. Plants of
the class marked by having two coty-
ledons; 27, 339, 340, 544.

Dicctyledonous (-eus). Having a pair of
cotyledons; 10, 314.

Didymous (-us). Twin, found in pairs.

Didyndmia. The Linnean class marked
by didunamy (335), 1. e.

Didynamous (-us). When a 4-androus
flower has the stamens in two pairs,
aud one pair shorter than the other;
250.

Dier¢silis. Mirbel’s name for a dry
fruit composed of several cells or car-
pels connate around a central axis,
and separating at maturity, as that of
Mallow.

Difformis. Of unusual formation.
Diffuse (-usus). Widely or loosely
spreading.
Digamous (-us).
same cluster.

Letting the light

Forked in pairs;

Of two sexes in the

407

Digitate (-atus). Fingered; a compound
leaf in which all the leaflets are borne
on the apex of the petiole; 101.

Digitately. In a digitate mode; same
as Palmately.

Digitate-Pinnate, 104.

Digynia. A Linnzean order character-
ized by having the gyneecium

Digynous. With two separate styles or
carpels; 261.

Dimerous (-us) Of two members in each
circle; 176.

Dimidiate (-atus). Halved, or as if one-
half was wanting.

Dimorphous (-us), Dimorphic, Dimor-

phism. Occurring under two forms;
225, 234.
Diecia. Linnean class (355) of plants

with the flowers

Diecicus (Dizcius, Dioicous). Unisex-
ual, and the two sexes borne by dis-
tinct individuals; 191.

Diecio-polygamous. When some indi-
viduals bear unisexual and others bi-
sexual flowers.

Dipetalous (-us). Two-petaled; 244.

Diphyllous (-us). Two-leaved; 243.

Diplo. See Duplo.

Diplostémonous, Diplostémeny. Having
twice as many stamens as petals or
sepals; 177, 198.

Diplotegium. <A capsule or other dry
fruit, invested with adnate calyx; an
inferior capsule.

Dipterous -(us). Two-winged.

Diremption (-io). Syn. of Chorisis; 202.

Disciferous (-us). Disk-bearing.

Disciform (-ormis). Depressed and cir-
cular, like a disk or quoit.

Discoidal or Discoid ( Discoideus). Ap-
pertaining toa disk. A discoid head
is one destitute of ray-flowers.

Disc or Disk (Discus). A word used in
several senses. The disk or dise of a
flower is a development of the torus
within the calyx, or within the corolla
and stamens; 213. In acapitulum or
head of flowers it is the central part
of the cluster, or the whole of it as
opposed to a border or ray. It is the
face or surface of any organ, such as
a leaf-blade, as opposed to the mar-
gin. In vegetable anatomy, certain
round spots or markings on cell-walls
are termed discs.

Discolor. When the two faces of a leaf,
&c., are unlike in color.

Discrete (-etus). Separate; not coales-
cent.

408

Disepalous (-us). Two-sepaled; 244.

Disk-jlowers. Those belonging to the
disk, or body, and not to the margin
or ray of a capitulum.

Dissected (-us). Deeply cut or divided
into numerous segments.

Dissépiment (-éntum). A partition in an
ovary or pericarp; 264.

Dissilient (-ens). Bursting asunder or
in pieces.

Distichous (-us}. Disposed in two ver-

, ticalranks; 122.

Distinct ( Distinctus).
not united.

Distractile (-ilis). Carried widely apart.

Diihécous (-us). Of two thecee, or cells,
as are most anthers; 254.

Diurnal. Daily ; occurring in the day;
sometimes used for ephemeral.

Divaricate (-atus). Extremely divergent.

Divergent, Diverging (-ens}. Inclining
away from each other.

Divided (Divisus). Where lobing or
segmentation extends to the base; 98.

Dodeca Greek for twelve. Used in

Dodecagynia. Linnan order with flow-
ers.

Dodecagynous. Having twelve styles or
distinct carpels.
Dodecamerous (-us).

the circle.

Dodecandria. A Linnean class (334)
with the flowers.

Dodecdndrous. Having twelve (or from
12 to 19) stamens; 249.

Dodrantalis. A span (about nine inches)
long.

Dolubriform (-ormis).
hatchet-shaped.

Dorsal (-alis). Relating to the dorsum
or back.

Dorsal Suture. That which answers to
the midrib of a carpel; 261.

Dorsiferous. Borne on the back.

Double. Has a technical use when a
flower is said to be ‘**double;”’ this
denoting one in which the leaves of
the flower are monstrously increased
mostly at the expense of the essential
organs.

Downy. Pubescent with fine nd soft
hairs. Loosely synonymous with soft-
pubescent, tomentose, Ke.

Drepanium. A sickle-shaped eyme; 156.

Drupaceous (-eus). Resembling or relat-
ing to a drupe.

Drupe (Drupa). A stone fruit; 297.

Drupelet, Drupel (Drupéola). A dimin-
utive drupe; 297.

Separate from;

Of twelve parts in

Axe-shaped or

GLOSSARY.

Drupetum. An aggregation of drupes;
300.

Duct. In vegetable anatomy, an elon-
gated cell or tubular vessel, ound espe-
cially in the woody (fibro-vascular)
parts of plants.

Dimetose (-osus). Pertaining to Dume-
tum, a thicket, or Dumus, a bush.

Dumose (-osus). Bushy, or relating to
bushes.

Duplo. Twice as many. In Greek com-
pounds, Diplo.

Duramen. ‘The heart-wood of an exo-
genous stem ; 80.

Dwarf. Of small size or height com-
pared with its relatives.

Dyclesium. See Diclesium.

E-or Ezx- As a prefix to Latin words,
carries a privative meaning, as Ecos-
tate, without ribs, Exalbuminous, with-
out albumen.

Eared. Same as Auriculate.

Ebracteate, Ebracteolate (-atus).
tute of bracts or bractlets.

Eburneus. Ivory-white.

Ecalcarate (-atus). Spurless.

Echinate (-atus). Beset with prickles,
like a hedgehog.

Echinulate (-atus).
tive prickles.

Edentate (-atus). Toothless.

Lffete (-etus), or Effetus. Past bearing;
functionless from age.

Efflorescence (-entia). The time or state
of blossoming; anthesis.

Effuse (-usus). Very loosely spreading,
more so than diffuse.

Eylandulose (-osus). Destitute of glands.

Eqgy-shaped. See Ovate.

Eladter. (ne of the spiral or spirally-
marked threads in the spore-cases of
certain Hepatice.

Eldtus. Tall or lofty.

Eleutheros. In Greek compounds, sep-
arate or distinct.

Eleutheropetalous (-us). Same as Chori-
petalous or Polypetalous; 245.

Ellipsoidal (-eus). An elliptical solid;
sometimes used for nearly elliptical.

Elliptical (-us). In the form of an el-
lipse. Oval or oblong with regularly
rounded ends: 95.

Emarcidus. Flaccid or withered.

Emarginate (-atus). With a notch cut
out of the margin; or, as usually ap-
plied, out of the extremity; 97.

Embracing. Clasping by the base.

Desti-

Beset with diminu-

GLOSSARY.

Embryo or Embryon. The rudimentary
plautlet formed in a seed; 9, 311.

Embiyonal. Relating to the embryo;
as Lmbryonal Vesicle ; 28+.

Embryo-sac. The cell in the ovule in
which the embryo is formed; 283.

Embryogeny. Embryo-formation.

Emersed (Emersus). Raised above and
out of the water.

Enantioblastus. With embryo at the end
of the seed diametrically opposite the
hilum.

£nation. Having outgrowths from the
surface, &e.; 179.

Endeca. In Greek compounds, eleven;
as in
Endecandrous, Endecayynous. With

eleven stamens or eleven styles, Ke.
Endemic. Contined geographically to
the particular region.
Endocarp (-drpium). The inner layer of
_a pericarp; 288
Endechrome (-oma). Peculiar coloring
matter in cells; especially the color-
, ing matter of Algw.
Endogens, Lndogene.
Plants; 70.
Endogenous structure, 70.
Endopleura. Inner seed-coat; 306.
Endophleum. Inner bark; 77.
Endorhizal (-us). Said of an embryo
which has the radicle sheathed by
the cotyledon or plumule wrapped
around it in many Monocotyledons ;
hence
Endorhize. Synonym of Monocotyle-
-, dones.
Endosperm (-ermium). Synonym of the
albumen of a seed; or the inner albu-
,men; 14, 310.
Endostome (-oma). The foramen of the
inner coat of an ovule; 277.
Endothecium. Inner lining of the cell of
an anther.
Enervis, Enervius.
or veins visible.

Endogenous

Nerveless; no ribs

Ennea. In Greek compounds, nine;
as in

Enneagynia. A Linnean ordinal name,

> and

Ennedgynous. With nine separate styles
or carpels; 337.

Enneandria. Linnean class, and Enne-
androus, with nine stamens; 249, 334.

Enédal (Enodis). Without a node.

Fnsatus. Same as Ensiform.

Ensiform (-ormis). Sword-shaped ; 7. e.
like a_broad sword, or the leaf of an
Iris. es?

409

Entire. Without toothing or division;
the margin whole and even; 97.

Entomophilous. said of flowers which
are habitually fecundated by pollen

, carried by insects ; 217, 218.

Entophytes (Entophyta). Plants grow-
ing in or out of other plants, as cer-
tain Fungi, &e.; 4.

Ephemeral. Lasting only for one day.

£pi. In Greek compounds, upon.

Lpiblast (-us). Name sometimes given
to the first (and an undeveloping) leaf
of the plumule of the embryo of grasses
and grain.

Lpiblastéma. A superficial outgrowth
from leaves, &c.; 210.

Epicalyz. Name sometimes given to
an involucel resembling an accessory

_ calyx.

Epicurp (Epicarpium). The external
layer of a pericarp; 288.

Epichilium. The terminal portion of
the labellum of an Orchid, when this
is of two parts.

Epiclinal (-us). Upon a torus.

Epicérolline. Upon a corolla.

Epidermis. The skin of a plant; 76, 89.

Epigeous (-eus) Growing on or out of
the ground.

Epigynous (-us). Literally on the pistil;
meaning on the ovary, or seemingly

185.

Epipetalous (-us). Borne on (adnate to)
the petals; also used in the sense of
piaced before the petals.

Epiphizum. The outermost or corky
bark; 76.

JF piphyllous (-us).~ Growing on leaves.

Fpiphytal. Pertaining to

Epiphytes (Epiphyta). Plants growing
on other plants by way of attachment,
but not parasitic; air-plants; 35.

Epipterous (-us). Winged or wing-
bearing at summit.

Episepalous. On the sepals; also used
in the sense of standing before a

_ sepal.

Episperm ( lpispermium).

_ outer coat of a seed; 305

Epitropors (-us). Name (by Agardh) of
an aratropous ovule with rhaphe
averse when ascending, adverse when
suspended: 282.

Equal (qualis). Alike as to length or

,number, &c., as the case may be.

Equitant '-ans). Riding: folded around,
as if straddling over; 108, 138:

Erect (£rectus). Standing upright,
mostly in relation to the ground,

sO;

The coat or

410

sometimes when perpendicular to the
surface of attachment.

Erion. Greek for wool; used in com-
pound words, as

Evianthus. With woolly flowers.

Erioéphorus. Wool-bearing.
Eriophyllus. Woolly-leaved.
Erostrate (Erostris). Beakless.

Erose (Lrosus). As if gnawed; applied
to an irregularly toothed or eroded
margin.

Erythros. Greek for red, used in com-
pound terms from the Greek.

Erythrostomum. Name given by Des-
vaux for such an aggregate fruit as
a raspberry; 300.

Estivation. See #stivation.
cterio. Name of aggregate fruits,

especially of fleshy ones, such as a
blackberry.

Etivlated. Blanched by darkness.

Eu. Prefixed to words of Greek deri-
vation denotes very, true, or much so.
Frequently used in names of sections
or other groups; 357.

Euphylia. Foliage-leaves, or true leaves.

Lutropic (-icus). Name suggested for
twining ‘‘with the sun;’’ dl.

Evalvular (Evalvis). Not opening by
valves.

Evergreen. Bearing green foliage all
the year rcund.

Evittatus. Not vittate.

Ex. A prefix in place of £ privative
when the following part of the com-
pound begins with a vowel; as

Exalbuminous (-osus). Destitute of albu-
men; 14, 309.

Exalate (-atus). Destitute of wing.
Ezxanthemata. Eruptive excrescenses on
the surface of leaves, &e.; blotches.
Exaristate (-atus). Destitute of an
arista or awn.
Exdsperate (-atus).

projecting points.

Excentric (-icus).
one-sided.

Excurrent (-ens). Running through to
the very summit or beyond; 48.

Exiguus. Small or mean.

Ezilis. Lank or meagre.

£zimivs. Distinguished, as for size or
beauty.

Fzo-. In Greek compounds, external or

, outward; as

Exocarp (Exocarpium). The outer layer
of a pericarp; 288.

Exé,enous. Outside growing. as the
wood of Dicotvledons; 69, 73.

Rough with hard

Out of the centre;

GLOSSARY.

Exogens, Exogene. Exogenous Plants;
69, 340.

Exorhize. Name equivalent to Exogen,
from

Ex hizal(-us). The radicle not sheathed,
so the primary root in germiration

, has no covering to break through.

Exostome (Exostoma). The foramen of
the outer coat of the ovule; 277.

Exothécium. The outer wall of an

, anther.

Explunate ( atus). Spread out flat.

Exsert, Exserted (Exsértus). Protruding
beyond or out of, as stamens beyond

_ the corolla.

Exstipulate (-atus). Destitute of stipules.

Exterior. External in the sense of outer.
But also in the flower sometimes used
in the sense of anterior.

FExtine. Outer coat of a pollen-grain.

Extra-azillary. Beyond or out of the axil.

Extrorse (Ezxtrorsus, Extrorsum). Di-
rected outward: 253.

Eye. A gardener’s name for an unde-
veloped bud.

Facies. Face; the general aspect.

Falcate (-atus), and Falciform (-ormis).
Scythe-shaped or sickle-shaped; plain
and curved, with the edges parallel.

Family. In botany, synonymous with
Order; 325.

Fan-shaped. See Flabelliform.

Farina. Starch.

Farinaceous (-eus). Of the nature of
starch, or containing it.

Farinose (-osus). Covered with a meal-
like powder.

Fasciate (-atus). Said of monstrous ex-
pansions of stems, giving the appear-
ance as of several stems coalescent in
one plane.

Fascicle (-icula). A close cluster or
bundle, whether of flowers, stalks,
roots, or leaves; 147, 153.

Fascicled (Fasciculatus, Fascicularis).
In a fascicle; 131.

Fastigiate (-atus). Said of branches
when parailel, clustered, and erect.
Faux, pl. fauces. The gorge or throat
of a gamophyllous calyx or corolla;
either at the orifice, or a portion
between the limb and the proper

tube; 246.

Faveolate (-atus), Favose (-osus). Honey-
combed; same as Alveolate.

Feather-veined. Waving veins all pro-
ceeding from the sides of a midrib.

Feathery. See Plumose.

GLOSSARY.

Fécula or Fecula. Starch-like matter.
Applied to a pistillate flower, or to a
pl nt producing only such flowers.

Fenistrate (-atus), Fenestralis. Pierced
with large holes, like windows.

Ferrugineous or Ferruginous (Ferru-
gineus). Colored to imitate iron-rust.

Fertile (-ilis). Fruitful, fruiting, or ca-
pable of producing fruit; as a fertile
flower is one provided with a well-
formed pistil; 191. In English deserip-
tions, Flower fertile usually means a
pistillate or female flower. Stamens
or anthers are also said to be fertile
when polliniferous and capable of fer-
tilizing.

Fertilization. Synonym of fecundation,
as of the ovule by pollen: 215.

Fibre (Fibra). Any fine filament; the
elementary components of wood, &c. ;
delicate roots, Ke.

Fibril (-illa) A diminutive fibre.

Fibrillate (-atus), Fibrillose (-osus).
Furnished or abounding with fibres
or fibrils.

Fibrous, Fibvose (-osus).
of the nature of fibres.

Fibro-vasculur. Consisting of woody
fibres and ducts.

Fiddle-shaped. Obovate and with a
sinus or contraction on each side.
Fidus. A Latin termination for cleft or

lobed.

Filament (-entum). The stalk or sup-
port of ananther; 165,251. Also any
fibre-shaped or thread-like body.

Filamentous, Filamentose (-osus).
posed of threads or filaments.

Filicology. The botany of Ferns: re-
placed by Pteridology.

Filiform (-ormis) . Thread-shaped; long,
slender, and terete.

Filipéndulous (-us).
thread,

Fimbria. A fringe, or dissected border.

Fimbriate (-atus). Fringed ; bordered
by slender processes or marginal ap-
pendages

Fimbrillate (-atus), Fimbriliferous (-us),
Bearing Fimbrille or diminutive fringe.

Fingered. See Digitate.

Fissiparous Multiplying by the divi-
sion of one body into two, and so on.

Fissus. Split or cleft. See Fidus,

Fistular, Fistulose (-osus). Hollow
through the whole length, as the leaf
and stem of an Onion.

Fldbellate (-atus), Flabelliform (-ormis).
Fan-shaped; much dilated from a

Composed or

Com-

Hanging from a

|

411

wedge-shaped base, and the broader
end rounded,

Flabellinerved. With radiating straight
nerves; 92.

Flaégellute (-atus), Flagellaris, Produe~
ing filiform runners (flagella), or
runner-like branches.

Flagelliform (-ormis). Runner-like;
long, slender, and supple like a whip-
lash or Flagellum ; 53.

Flammeus. \lame-colored.

Flavescent (-ens). Yellowish or pale
yellow.

Flavus. Pale yellow or ochre-yellow.

Fleshy. Succulent; of the consistence
of flesh.

Flexuous, Flexuose (-vsus). Zigzag;
bent alternately im opposite direc-
tions.

Floating. Borne on the surface of water.

Floccove (-osus). Bearing or clothed with
locks of soft hairs or wool ( flocc7).

Flocculent. Diminutive of floccose.

Flora (Goddess of flowers). The aggre-
gate of the plants of a country or dis-
trict; or the name of a work which
systematically describes them; 369.

Floral. Belonging to the flower.

Floral Envelopes. YFlower-leaves; 164.

Floret. A small flower, one of a cluster.

Flovibundus. Abundantly floriferous.

Flovifcrous( Florifer, Floriferus). Bear-
ing flowers

Flérula. A small Flora; the Flora of
a restricted district.

Flos. Latin for flower. — Flos plenus. A
“ double’? flower; that is, one in which
petals are increased abnormally, com-
monly at the expense of the andree-
cium or the gyneecium also; 171.

Floésculus. Latin for floret.

Flower. The whole reproductive appa-
ratus in a phenogamous plant; 163.
Flower-bud. An unexpanded blossom

or undeveloped cluster; 40.

Flowering Plans, 3, 344.

Flowerless Plints, 3, 344.

Fluitans. Floating.

Fliviatile, Fluviatilis. Belonging to a
river or running water.

Fly-traps, 113

Femineus. Feminine or female flower,
plant, &e.; 191.

Foliaceous (-eus). Leaf-like in texture
or appearance; or bearing leaves.

Foliar (Foliaris). Relating to leaves.

Foliation (Feliatio). WLeafing out.

Foliate (-atus). Waving leaves. With
Latin numerical prefix, 5/foliale, tri-

412

foliate, and so on, according to the
number.

Foliijvrm (-ormis). Leaf-shaped.

Foliolate (-atus). Having leaflets: their
number may be indicated by Latin
numerals, as b/foliolate, trifoliolate,
&e.; 102.

Foliilum. <A Jeaflet; 102.

Folivse. Bearing numerous leaves.
Folium. Latin tor leaf; 85.

Follicetum. A whorl or aggregation of
follicles; 300.

Follicle (-iculus). Fruit of asingle carpel
dehiscent by one (the ventral) suture;
292. Name of the earlier botanists
for any kind of capsular fruit.

Follicular (-aris). Pertaining to or like

- a follicle.

Foot-stalk. Petivle, 85; or Peduncle, 143.

Fordmen. An aperture of any kind;
specially that of the coat of the ovule;
277.

Foraminulose (-osus). Pierced with many
small holes.
Forcipate (-atus).

cers.

Forked. Divided equally into branches.

Fornicate (-atus). Arched over, as by
scales (Fornices) covering the throat
of the corolla of Hound’s-tongue, &e.

Foveate (-atus), and diminutive Foveo-
late. Pitted; impressed with shallow
depressions or pits, Fovee.

Foville. Minute granules in a liquid,
in the protoplasm of the pollen-grain,
&e.; 258.

Free. Not adnate to other organs.
Some imes used in the sense of dis-
tinct, 7. e. unconnected with others of
the same sort.

Fringed. See Fimbriate.

Frond (Frons). An old name for leaf:
employed mainly for the leaf of Ferns
and other Cryptogamia, and certain
Phenogamous plants which serve for
fructification as well as foliage; also
for the peculiar foliage of Palms; 67.

Frondescence (-entia). The act of leaf-
ing. Has also been employed to
express the metamorphosis of floral
organs into foliage-leaves; 174.

Frondose (-osus). Sometimes used in
the sense of leafy; also frond-like, or
bearing fronds.

Fructification. The act or the organs
of fruiting or reproduction through
flower and seed, or their analogues.

Fructus. Latin for Fruit.

Fruit. The immediate product of fruc-

Like forceps or pin-

GLOSSARY.

tification ; in phenogamous plants.
the seed-vessel and contents, along
with all intimately connected acces-
sory parts; 285.

Fruit-dots in Ferns. See Sorus.

Frumentaceous. Relating to grain (Fru-
mentum).

Frustulose (-osus). Consisting of similar
pieces or Frustules (frustula).

Frutex. A shrub.
Frutescent (-ens). Shrubby, or becom-
ing so.

Fruticose (-osus).
to shrubs.

Fruticulose (-osus). Relating to a di-
minutive shrubby plant.

Fruticulus, A minute or low shrubby
plant.

Fugacious. Falling or fading very early;
lasting a very short time.

Fulvous (-us). Tawny; orange-yellow
and gray mixed.
Fulcra. Accessory organs, such as ten-
drils, stipules, spines, and the like.
Fulcrate (-atus). Propped, supported
by, or provided with accessory organs.

Fuliginous (-osus). Sooty-brown.

Fungiform (-ormis and Fungilliformis).
Mushroom-shaped.

Fungose (-osus). Spongy in texture;
fungus-like.

Finicule, Funiculus.
ovule or seed; 276.

Funnelform, Funnel-shaped; 249.
Infundibuliform.

Furcate (-atus). Forked; or divergently
branched.

Furcellatus. Diminutively forked.

Furfuraceous (-eus). Scurfy; covered
with bran-like scales or powder.

Furrowed. See Sulcate.

Fuscous (-us). Grayish-brown in hue.

Fusiform (-ormis). Spindle-shaped ;
terete and tapering gradually to each
end; 31.

Shrubby, or relating

The stalk of an

See

Gdlbulus. The peculiar strobile of Cy-
press and Juniper, composed of up-
wardly thickened or fleshy scales;
393.

Galea. A helmet; name given, from
its shape, to the upper sepal of Aconi-
tum, and the upper lip of certain
forms of bilabiate corolla; 247.

Galeate (-atus): Having a galea; hel-
met-shaped; 247.

Gumo-. In Greek compounds, denotes
union by the edges or coalescence.

GLOSSARY.

Gamopetalous. A corolla of coalescent
petals ; formerly Monopetalous ; 244.

Gamophyllous (-us). Composed of coa-
lescent leaves.

Gamosepualous.
sepals; 244.

Geitondgamy. Fecundation of a pistil
by pollen of another flower of the
same plant; 216.

Geminate (-atus).
side by side.

Gemma. A bud, specially a leaf-bud.

Gemmation. Budding-growth; or the
disposition of buds.

Gemmule (Gémmula). Diminutive of
gemma; minute and simple buds or
bodies analogous to buds; also sy-
nonym of Plumule. Fora time used by
Endlicher and others for the ovule.

Genera. Plural of Genus; 323.

General (-alis). Opposed to partial; as
general inyolucre.

Generic, Relating to genus.

Genetic. Genealogical ; that which comes
by inheritance.
Geniculate (-atus).

a knee.

Genitalia. The stamens and pistils or
their analogues.

Genus. Kind or group superior to spe-
cies, and which with the species gives
the name to the plant; 323.

Geoblast (-astus). A plumule which in
germination rises from underground,
such as that of the Pea.

Germ. A growing point or initial growth,
as of a bud; or the Embryo; 311. Or
in the sense of

Germen. The Linnean name of the
ovary; 166. :

Germination (-atio). The act of devel-
opment of the embryo of a seed into a
plant.

Gerontogeous (-eus).
Old World.

Gibbous, Gibbose (-osus) Swelling out
on one side into a gibber or gibber-
osity.

Gigantéus. Of unusual height.

Gilrus.. Dirty yellow with a tinge of red.

Glabrous ( Glaber). Smooth in the sense
of not pubescent or hairy.

Glabrate (-atus). Somewhat glabrous,
or becoming glabrous.

Glabriisculus. Almost but not quite
glabrous.

Gladiate (-atus). Sword-shaped ; in the
form of a sword-blade, whether straight
or somewhat curved. See Ensiform.

A calyx of coalescent

Twin; in pairs; two

Bent abruptly, like

Belonging to the

413

Gland for Glans. An acorn and the
like; 296.

Gland (Glandula). A definite secreting
surface or structure on the surface of
any part of a plant, or partly imbedded

" in it, extended to any protuberance
or structure of similar nature which
may not secrete.

Glandular, Glandulose (-osus). Bearing
glands or having the nature of glands.

Glanduliferous (-us). Gland-bearing.

Glareosus. Growing in gravel.

Glaucescent (-ens). Verging upon or be-
coming glaucous.

Glaucous (-us). Covered or whitened
with a bloom, like that on a Cabbage-
leaf.

Globose (-osus). Having or approaching
a spherical form.

Globular (-aris), Globulose (-osus). Some-
what or nearly globose.

Glochideous, Glochidiate (-atus). When
bristles and the like are barbed at tip.

Glochis. A barb.

Glomerate (-atus). Compactly clustered,
especially into a

Glémerule (Glomérulus, Glomus). <A
cyme condensed into a head or capi-
tate cluster; 152.

Glossology, 3, 358.

Glumaceous (-€us).
sembling glumes.

Glume, Gluma. One of the chaff-like
bracts of the inflorescence of Grasses
and their relatives; 143.

Glumella. Diminutive of gluma; an
inner or secondary glume.

Glutinous (-osus), Covered with a sticky
exudation.

Gonophore (Gonéphorum). A stipe which
elevates both stamens and pistil; 212.

Gossypine (-inus). Cottony; flocculent.

Gracilis. Slender.

Grain. See Caryopsis.

Gramineous (-eus). Relating to grass or
grain-bearing plants.

Granular (-aris), Granulose (-osus).
Composed of small grains or Granulcs.

Granulate (-atus), Granuliferus. Bear-
ing grains or grain-like bodies.

Gravéolens. Unpleasantlystrong-scented.

Griseus. Gray or bluish-gray.

Grumous (Grumosus). Consisting of
clustered grains.

Guttate (-atus). Spotted as if by drops.

Gymnanthous (-us). Naked flowered.

Gymnes. Greek for naked; used in com-
pounds such as

Gymnocarpous (-us).

Pertaining to or re-

Naked-fruited.

414

Gymnospermia. A Linnean artificial
order of Didynamia, in which the
nutlets resulting from four divisions
of an ovary were taken for naked
seeds; 337.

Gymnesperms, Gymnosperme. A sub-
class of naked-seeded plants; 268, 344.

Gymnospermous (-us). Naked-seeded, as
opposed to Angiospermous.

Gynandria. A Linnean class, character-
ized by the flower being

Gynandrous. Stamens borne on (adnate
to) the pistil, even to the style or
stigma; 251, 335.

Gynobase (Gynobasis). An enlargemeut
or production of the torus on which
the gynecium rests or is somewhat
elevated; 212.

Gyno-diecious. _ Dicecious with some
flowers hermaphrodite and others pis-
tillate only; 191.

Gynacium. The pistil or collective pis-
tils of a flower; the female portion of
a flower as a whole; 165.

Gynophore ( Gynéphorum).
a pistil; 212.

Gynostegium. A sheath or covering of
the gvneecium, of whatever nature.

Gynostemium. The column of an Orchid,
consisting of andreecium and summit
of the gvneecium combined.

Gyrate (-atus). Curved into a circle, or
taking a circular course.

Gyrose (-osus). Curved backward and
forward in turus.

The stipe of

Habit (Habitus).
ance of a plant.

Habitat. Habitation; the geographical
limits or station; 366.

Hematitic (-icus). Brown-red.

Hairs. Outgrowths of the epidermis,
consisting of single elongated cells, or
of a row of cells.

Hairy. Descriptively applied to pilosity
or pubescence, in which the hairs are
separately distinguishable.

Hulbert- or Halberd-shaped.
tate.

Halved. See Diinidiate; with one half
absent or appearing to be so.

Humate (-atus). Hooked at the tip.

Hamulate or Hamulose (-osus). Dimin-
utive of Hamate.

Haplos. In Greek compounds, simple
or simply, as

Haplopetalus (-us).
of petals.

The general appear-

See Has-

With only one row

}

GLOSSARY.

Haplostemonous (-us). With .a single
series of stamens; 177.

Hastate (-alus), Hastilis. Halberd-
shaped, like the head of a halberd,
v7. €. sagittate, but the basal lobes di-
rected outward or at right angles to
the midrib of the leaf; 96.

Head. The form of inflorescence termed
Capitulum, viz. a cluster of sessile
flowers ona very short axis and centvi-
petal in evolution; 147.

Heart-shaped. Ovate with a sinus at
base; 96.

Heart-wood. The older and matured
wood of an exogenous stem; 80.

Hcbetate (-atus). Having a dull or blunt
and soft point.

felicoid (-oideus), Helicoidal. Coiled into
a helix, or like a snail-shell. In true
helicoid inflorescence, the flowers are
all in a single row; 155, 157.

Helmet. See Galea.

Helwolus. Dull and grayish yellow.

Hemi. Walf or halved; in Greek com-
pounds, such as

Hemi-anatropous. Half anatropous.

Hemicarp (-arpium). Half or one carpel
of a Cremocarp.

Hemitropous (-us). Same as amphitro-
pous or half anatropous.

Hepta, The Greck numeral seven.

Heptagynia. A Linnean artificial order,
having seven styles or distinct car-
pels; 337. ;

Heptamerous (-us). Of seven members.

Heptandria. The Linnzan class with
seven stamens; 334. :

Heptandrous. Seven-stamened; 249,

Herb (Herba). A plant with no persist-
ent woody stem above ground; 50.

Herbaceous. Of the texture, color, or
other characters of an herb.

Herbarium, Herbal. A collection of
dried specimens of plants, systemati-
cally arranged; 380.

Hercogamous (-us). Said of hermaphro-
dite flowers when some structural ob-
stacle prevents autogamy.

Hermaphrodite (-itus). Of both sexes;
191.

Hesperidium. A hard-rinded berry, like
an orange and lemon; 299.

Heteracmy. Synonym of Dichogamy ;
219
Heteros. In Greek compounds, denotes

(liverse or various, as

Heterocarpous (-us, Heterocarpicus..
Producing more than one kind of
fruit.

GLOSSARY.

Heterocephalous (-us). Bearing two
kinds of head or capitulum.

Tleteroclite (-itus). Anomalous in forma-
tion,

Ilctcrocline (-inus). Nearly same as
Heterocephalous ; on separate recepta-
cles.

Heterodromous (-us). Spirals of changing
direction.

Heterogamous (-us). Bearing two kinds
of flowers.

Heterogeneous. Not uniform in kind.

Heteroyone or Heteroyoneus. When the
flowers are dimorphous or trimor-
phous as respects relative length, &c.,
of stamens and pistil; 225, 234.

Heterostyled. Same as Heterogone ; 234.

Heteromerous. Of members not corre-
sponding in number.

Heteroplhyllous (-us). Having leaves of
more than one form.

Heterotropous (-us). Turned in more
than one direction, or in an unusual di-
rection; same as Amphitropous; 279.
(Also used by Agardh for collateral
ovules turned back to back; 282.)

Heza. Greek numeral six; from which
is formed

Hexagynia. Linnean artificial order, of
flowers with six styles or distinct car-
pels; 337.

Hexayynous.
Hexagynia.

Hexamerous (-us). Of six members; 176.

Hexandria. Linnean class with perfect
flowers of six stamens; 334.

Hexandrous. Having six stamens; 249.

Hezxapetalous (-us). Having six petals.

Hexaphyllous. Six-leaved.

Hexapterous (-us) Six-winged.

Hexasepalous. Waving six sepals.

Hexastémonous. Having six stamens.

Hibernaculum. A winter-bud; 40.

Hidden. Concealed from sight; as

Hidden-veined, where the veins are in-
visible, as in the leaves of Pinks and
Houseleeks.

Hiemal (-alis). Relating to winter

Hilar ( Hilaris). Belonging to the hilum.

Hilum. The scar or place of attachment
of the seed; 277, 305.

Hippocrepiform (-ormis).
shaped.

Hirsute (-utus). Pubescent with rather
coarse or stiff hairs.

Hirtellous (-us). Minutely hirsute.

Hirtus. Hairy, nearly same as Hirsute.

Hispid (-idus). Beset with rigid or
bristly hairs or with briztles.

Having the character of

Horseshoe-

415

Hispidulous (-us). Minutely hispid.

Hoary. Grayish-white with a fine and
close pubescence. See Canescent.

Holosericeous (-eus). Covered with fine
and silky pubescence.

Homocarpous(-us). With fruit all of one
kind.

Homodromous (-us), Homodromy.
spirals all of uniform direction.

Homogamous (-us). Bearing one kind
of flowers.

Homogeneous. All of one nature or kind.

Homégonous or Homogone. Homomor-
phous as respects the stamens and pis-
til; opposed to dimorphous; 225.

Homologue. A homologous organ or
part.

Homologous. Of one name or type, such
as leaves and parts answering morpho-
logically to leaves; 6.

Homomdilus. Said of leaves and the like
which are all turned in one direction.

Homomorphous (-us). All of one form.

Homostyled. Same as Homogone.

Homotropous (-us). Curved or turned in
one direction; applied also to the em-
bryo of an anatropous seed, with rad-
icle next the hilum; 312.

Wood. See Cucullus.

Hooded, Bearing or in form of a hood.

Hornus, Hornotinus. Of the present
year.

Horny. Of the consistence of horn. See
Corneus.

Hortensis, Hortulanus.
the garden.

Hortus siccus. Old name of an herbarium.

Humi. On the ground.

Humifisus, Humistratus.
the surface of the ground.

Humilis. Low of stature.

Hyaline (-inus). Transparent or trans-
lucent.

Hybrid. A mongrel, or cross-breed of
two species; 321.

Hydrophytes (Hydrophyta).
plants.

Hyemalis. See Hiemalis.

Hypanthium. An enlargement or other
development of the torus under the
calyx; 214.

Hypo. In Greek compounds, denotes
under, beneath, lower.

Hypanthodium. Same as Syconium;
149, 303.

Hypochilium. The basal portion of the
labellum of an Orchid.

Hypocraterimorphous (-us), or Hypoera-
teriform, but the latter is a hybrid of

With

Pertaining to

Spread over

Water-

+16

Greek and Latin. Salverform or sal-
ver-shaped; that is, in the form of a
salver raised on a central support or
stem beneath. Said of a corolla and
the like with slender tube abruptly ex-
panded into a flat limb; 248.

Hiypogeous (-eus). Growing or remain-
ing underground; 19.

Hypogynous (-us). Under or free from
the gyneecium or pistil; 182.

Hypophyllous (-us). Growing on the un-
der side of a leaf.

Hypophyllium. An abortive leat or scale
under another leaf, or seeming leaf, as
in Asparagus and Ruscus.

Hypsophylla. Answers to the German
‘*Hochblatter,” or high leaves, those
of the inflorescence, i. e. bracts and
the like; 6.

Hysteranthous (-us). With leaves pro-
duced later than the blossoms.

Icosandria. The Linnean class with
twenty stamens (as the name denotes)
or a larger number, inserted on the
calyx; 334. /cosandrous is the corre-
sponding adjective; 249.

Imberbis. Not bearded. :

Imbricate (-atus), Imbricative. Over-
lapping so as to ‘break joints,”’ like
tiles or shingles on a roof; either with
parts all in one horizontal row or cir-
cle, as in the wstivation of a calyx or
corolla, when at least one piece must
be wholly external and one internal;
or with the tips of lower parts covering
the bases of higher ones in a succession
of rows or spiral ranks; 135.

Immarginate (-atus). Not margined or
bordered.

Immersed (-us).
water.

JImpari-pinnate. Pinnate with an odd
terminal leaflet; 101.

Inequilateral (-alis). Unequal-sided.

Inanis. Empty, as an anther containing
no pollen.

Inappendiculate (-atus).

Growing wholly under

Not appen-

daged.

Incanescent. Same as Canescent.
Incanus Hoary-white.

Incarnate (-atus). Flesh-colored. See
Carneus.

Incised (-us). Cut irregularly and

sharply; 98.

Included (Inclusus). When the part in
question does not protrude beyond the
surrounding organ.

GLOSSARY.

Incomplete (-us). Wanting some essen-
tial component part; 190.

Incrassate (-atus). Thickened.

Incubous (-us). The tip of one leaf or
other part lying flat over the base of
the next above it.

Incumbent (-ens).
upon.

Incumbent Anther. One lying against
the inner face of filament; 253.

Incumbent Cotyledons, when the back
of o:e lies against the radicle; 313.

Jncurved (-us). Bending from without
inward.

Indefinite (-itus). Relates usually to
number, this either uncertain or too
many for easy counting.

Indefinite Growth, 49.

Indefinite Inflorescence, same as Inde-
terminate; 144. ;

/ndehiscent (-ens). Not opening by valves,
chinks, or along regular lines; 288.

Indeterminate. Not terminated abso-
lutely, as the inflorescence in which
no blossom ends the axis of the flower-
cluster; 144, 146.

Indigenous (-us).
to the country.

Individuals, 315.

Leaning or resting

Native and original

Indivisus. Undivided, i. e. not cleft,
lobed, or parted.
Indumentum. Any hairy covering or

pubescence which forms a coating.

Induplicate (-atus). With edges folded
in or turned inward.

Indusium. The proper (often shield-
shaped) covering of the sorus or fruit-
cluster of a Fern.

Induviate (-atus). Clothed with with-
ered parts or /nduvie (clothing).

Inequilateral. Unequal-sided; 106.

Jnermis. Unarmed, without prickles,
thorns, &e.

Inferior (Inferus). Said of one organ
when below another. In the blossom
also in the sense of anterior; 160. An
inferior calyx is one below the ovary,
or free; 183. An inferior ovary is one
with adnate or superior calyx ; 183.

Inflated (-atus). Bladdery.

Inflexed (-us). Bent or turned abruptly
inward.

Inflorescence. Mode of disposition of
flowers ; less properly used fora flower-
cluster itself; 141.

Infra-azillary (-aris). Below the axil.

Infundibuliform (-ormis), Infundibular
(-aris). Funnelform, funnel-shaped;
249.

GLOSSARY.

Innate (-atus). Borne on the apex of
the supporting part; in an anther the
counterpart of adnate; 252.

Innovation (-i0). A new-formed shoot.

Inosculating. Same as Anastomosing.

Inseparvate, Inseparation. Terms pro-
posed by Masters to express coales-
cence; 181.

Inserted (-us).
out of.

Insertion (-io0). Is the mode or place
where one body is attached to that
which bears it.

Integer. Entire in the sense of un-
divided, or not lobed ; 97.

Jategérrimus. Entire in the sense of
quite entire, z.e. the margin without
dentation; 97.

Inter. Between; as in Jntercellular,

_ between the cells, &e.
Interfoliaceous (=eus).

Attached to or growing

Between the

leaves of a pair, as the stipules of |

many Rubiacee.
Internode (-odium). The portion of
stem between two nodes; 6.
Interpetiolar (-avis). Between the pet-
ioles.
Interruptedly pinnate.
out a terminal leaflet.
Intine. The inner coat of a grain of
pollen.
Intrafoliaceous (-eus).
fure a leaf.
Introflezed (-us). Same as inflexed.
JIntrorse (-orsus). Turned inward or
toward the axis; 253.

Pinnate with-

Within or be-

Introvenius. Same as hidden-veined.
Intruse (-usus). Pushed or projecting
inward.

Involicellate (-atus). Provided with a
secondary involucre or

Fnvolucel (-ellum). An inner or secon-
dary involucre that of an umbellet,
&e.; 142.

Involicrate (-atus).
involucre.

Involucre (Involic: um). <A circle of
bracts subtending a flower-cluster;
142.

Involute (-utus). Rolled inward: 133.

Irregular (-aris). Exhibiting a want of
symmetry in form; 184.

Irregularity, 179, 184, 219.

Isadelphous (-us). Equal brotherhood, as
when the number of stamens in two
phalanges is equal.

Iséchrous. All of one color or hue.

Isémerous (-us). The members of suc-
cessive circles equal in number; 175.

Provided with an

27

417

Isostémonous (-us). The stamens just as
many as the petals, &e. ; 177.
Tsustemony, 196.

Jointed. See Articulated.

Juba. A loose panicle, with axis de-
liquescent.

Jugum, pl. juga. <A pair of leaflets. So
pinnate leaves are unijugate, with a
single pair of leaflets; bijugate, with
two pairs; tryugate, with three pairs
or juga, &c. Also the ridges on the
fruit of Umbeiliferze are termed juga.

Julus. Same as Amentum or Catkin.

Julaceous (-eus). _Catkin-like, Amen-
taceous.

Keel. A central dorsal ridge, like the
keel of a boat. The two anterior
petals of a papilionaceous corvlla,
which are united into a body shaped
like the keel or the prow of a vessel;
185.

Keeled. Having a keel. See Carinate.

Kernel. The nucleus of an ovule, or
of a seed, z. e. the whole body within
the coats.

Kermesinus. Of the color of carmine.

Key-fruit. See Samara; 294.

Kidney-shaped. Crescentic with the
ends rounded; very oblately cordate;
96.

Kingdom, 325.

Labellum. One of the petals of an Or-
chideous flower, which is unlike the
others.

Labiate (-us).
247.

Labiatiflorous (-us). Said of certain
Composite with bilabiate corollas.

Labiose (-osus). Said of a polypetalous
corolla which has the appearance of
bilabiation.

Labium. See Lip.

Lacerate (Lacerus).
if torn or lacerated.

Lacinia. A slash; used for a slender
lobe.

Laciniate (-atus).
narrow incisions.

Lacinula. A diminutive lacinia or nar-
row lobe.

Lactescent (-ens). Yielding milky juice.

Lacteus. Milk-white.

Lacunose (-osus). Abounding in pits,
holes, or depressions (/acune).

Lacistrine (Lacustris). Belonging to or
living in lakes or ponds.

Lipped, mostly Bilabiate;

Irregular cleft as

Slashed; cut into

418

Levigate (-atus). Smooth as if polished.

Levis (this form, and not levis, has al-
ways been used in botany). Smooth
in the sense of not rough.

Lageniform (-ormis). Shaped like a
florence flask or a gourd (the fruit of
Lagenaria).

Layopus. Hare-footed.
ered with long hairs.

Lamella. A thin plate.

Lamellar (-aris), Lamellate (-atus), La-
mellose (-vsus). Composed of thin
plates or lamellz.

Lamina. The blade or expanded part
of a leaf, &e ; 85, 245.

Lanate (-atus), Lanose (-osus). Bearing
long and implexed hairs or wool
(lana).

Lancevlate (-atus). Shaped like a lance
or spear-head; narrower than oblong,
and tapering to each end, or at least
to the apex; 95.

Lanuginous (-vsus). Cottony or woolly ;
clothed with soft aud implexed hairs
or down (lanigo).

Densely cov-

Lappaceus. Like a bur (lappa).
Lasianthus. Woolly-flowered.
Latent. Undeveloped or dormant, as

certain buds; 40.
Lateral (-alis). Belonging to or borne
on the sides.

Lateritious (-ius). Of a brick-red color.

Latex. Proper juice, milky juice, and
the like.

Laticiferous. Containing or conveying
latex.

Latiseptus. With broad partition.

Lavender-color. Pale blue with some
gray.

Laxus. Loose.

Leaf. The principal sort of appendage
or lateral organ borne by the stem or
axis; 85.

Leaf-blade. The lamina of a leaf.

Leaf-bud. A bud which develops into
a leafy branch or its continuation; 40.

Leaflet. A blade or separate division
of a compound leaf; 100.

Leaf-scar. The cicatrix left by the ar-
ticulation and fall of a leaf; 47.

Leafstalk. -A petiole or footstalk to a
leaf-blade; 85, 104.

Leathery. See Coriaceous.

Lecus. A synonym for Corm.

Légume (Legimen). The seed-vessel of
Leguminose, a carpel which normally
dehisces by both the ventral and the
dorsal suture; 292.

Leguminous. Pertaining to a legume,

GLOSSARY.

or to the order to which the legume
gives its name.

Lenticels (Lenticélle).
on young bark.

Lenticular (-aris). Lens-shaped, that is
the shape of a lentil or a double-convex
lens.

Lentiginosus. Covered with minute dots
or freckles.

Lepal, Lepalum. A made-up word to.
signify a stamen transformed into a
scale, nectary, &e.

Lepicéna. Unused name for a glume of
Grasses.

Lepidote (-otus). Beset with small scurfy
scales.

Lepis. Greek term for a scale.

Leptos. Greek for slender; as in Lep-
tophyllus, slender-leaved.

Leucos. Greek for white; whence

Leucanthus. | White-flowered.

Leucophyllus. White-leaved, &e.

Liber. The inner and often fibrous
bark; 77, 81,

Lid. See Opereulum.

Ligneous (-eus), Lignosus. Woody.

Ligule (Ligula). A strap or strap-
shaped body, such the principal part
of a ray corolla in Composite. The
thin and scarious projection from the
summit of the sheath of the leaf of
Grasses, &c.; 106. Or a similar out-
growth of the inner face of certain
petals; 211.

Ligulate (-atus), Liguliform (-ormis).
Furnished with a ligule; 148, 247.
Liguliflorous (-us). Said of the head of
those Composite which contain only

ligulate corol/as.

Liliaceous. Lily-like; 246.

Limb (Limbus). A border, 7. e. the ex-
panded part of a gamophyllous peri-
anth, &c., as distinguished from the
tube and threat; 245. Sometimes the
term is applied to the lamina or blade
of a petal or a leaf.

Limbate (-atus). Bordered.

Line (Linea). The twelfth part of an
inch. By some reduced to the tenth
of an inch; but the decimal line is un-
usual in botanical measurement.

Linear (-aris). Narrow, several times
narrower than wide, and the margins
parallel; 95.

Lineate (-atus).

Lineolate (-atus).
obscure lines.

Lingueformis or Linguiformis, also Lin-
culate (-atus). ‘Tongue-shaped.

Lenticular spots

Marked with lines.
Marked with fine or

GLOSSARY.

Lip. One of the two divisions of a bila-
biate corolla or calyx, 1. e. of a gamo-
phyllous organ which is cleft into an
upper (superior or posterior) and a
lower (inferior or anterior) portion or
lip (labium).

Litoral or Littoral (-alis). Belonging
to or growing on the seashore or river-
shore.

Livid (-idus). Pale lead-colored.

Lobe (Lobus). Any division of an organ;
or specially a rounded division or pro-
jection; 98.

Lobate (-atus) or Lobed. Divided into
or bearing lobes; 98.

Lobulate (-atus). Divided into small
lobes, Lobdelets, or Lobules; 98.

Lécellate (-atus). Divided into locelli;
263.

Locellus. A secondary cell; as where
a proper cell (/oculus) of an anther or
an ovary is divided by a partition into
two cavities; 251, 263.

Léculament (-entum). Sume as Loculus;
289.

Locular (-aris). Celied; as Dilocular,
two-celled; ¢trilocular, three-celled,
quadrilocular, four-celled, &c.

Loculicidal (-idus), Loculicide. Dehis-
cent into the cell or cavity of a peri-
carp by the back, 7. e. through a dor-
sal suture; 289.

Léculus. The cell or cavity in an ovary
or an anther.
Loculose (-osus).

cells.

Lécusta. Name of a spikelet in Grasses.

Loédicule, Lodicula. One of the small

- seales next to the stamens in the flower
of Grasses.

Lomentaceous (-eus).
sembling a

Loment (Lomeéntum). A legume which
is constricted or which separates into
one-seeded articulations; 293.

Lorate (-atus). Strap-shaped or thong-
shaped; same as much-elongated
linear. :

Lucid (-idus). With a shining surface.

Lunate (-atus). Half moon-shaped;
crescent-shaped.

Linulate (-atus). Diminutive of Lunate.

Lipuline (-inus). Resembling a head of

Partitioned off into

Bearing or re-

Hops.

Lurid (-idus): Dingy-brown.

Lusus. A ‘sport’? or variation from
seed or bud: 319.

Lutéolus. Yellowish; diminutive of
Luteus.

41%.

Lutescent (-ens).
faintly yellow.

Liteus. Latin for yellow.

Lycotropous (-us). Said of an ortho-
tropous ovule when bent into an open
curve or horseshoe form.

Lyrate (-atus). Lyre-shaped; a pinnat-
ifid form with terminal lobe large and
rounded and one or more of the lower
pairs small; hence Lyrately pinnate ;
101, &e.

Becoming yellow, or

Macros. Greek for large or more prop-
erly long; hence Macranthus, long-
flowered; | Macrocephalus, large-
headed; Macropodus, long-footed, or
with long stalk, &e.

Macrospore. The larger kind of spore
in Lycopodiacex, &c.

Maculate (-atus). Spotted or blotched,
i.e. with macule.

Malpighiaceous Hairs. Those fixed by
the middle and tapering both ways.
Mamillate (-atus), Mamillar (-aris).

Bearing teat-shaped jfrocesses.

Mammeform (-ormis). Breast-shaped

or teat-shaped; conical with rounded

apex.
Mammosus. Breast-shaped.
Mancus. Deficient or wanting.

Manicate (-atus). Said of pubescence
so dense and interwoven that it may
be stripped off like a sleeve.

Marcescent (-ens), Marcidus. Withering
without falling off; 243.

Maryginate (-atus). Furnished with a
margin of a distinct character or ap-
pearance.

Marginicidal. Dehiscent by the dis-
junction of the united margins of car-
pels; 290.

Marmoratus. Marbled;- traversed by
veins or shades of color.

Maritime (Maritimus). Pertaining to
the sea or seacoast.

Mas, Masculus, Masculinus. Belonging
to the stamens, or staminate plant, or
flower; 191, &e.

Masked. Sec Personate.

Mealy. See Farinaceous.

Medial, Median (Medianus). Belong-
ing to the middle; in the plane of
bract and axis; 160.

Medificus. Fixed by the middle; 253.

Medilla. Pith; 75.

Medullary. Relating tothe pith J/ed-
ullary Rays, 74; Medullary sheath, 75,

Meiostémonous. With fewer stamens
than petals.

420

Melleus. Having the taste or smell of
honey.
Melligo. Honey-dew.

Membranous, Membranaceous (-eus).
Thin and rather soft or pliable, like
a membrane.

Meniscoid (-oideus).
like a meniscus.

Mericarp (-arpium). One of the akene-

Concayo-conyex,

like carpels or a closed half-fruit of |

Umbellifera ; 297.
Mevismatic. Dividing into parts or sim-
ilar portions.
Merithallus. Synonym of Internode.
Merous (-us). In Greek compounds, de-
notes parts or members; lence Dime-
rous, of two parts, &e.
Mesocarp (-arpium). The middle layer
of a pericarp; 285.
Mesophleaum. ‘The middle or green
bark ; 76.
Metamorphosis, Metamorplhy, 167.
Micropyle (-yla). The spot or point in
the seed at which was the orifice of the
ovule; 277, 305.
Microspore. The smaller kind of spore
in Lycopodiacee, &c.
Midrib. The central or main rib; 92.
Miniate (-atus). Vermilion-color.
Mistus or Miztus. A cross-breed ; 321.
Mitreform or Mitriform (-ormis). Mi-
tre-shaped or cap-shaped.
Monadelphia. The Linnzean class con-
taining flowers with Monadelphous
stamens, 7. €. those united by their
filaments into a tube or column; 250,
335.
Monandria. The Linnzan class (334)
“containing flowers with Jfonandrous,
that is, a single stamen; 249.
Monadnthous (-us). One-flowered.
Moniliform (-ormis). Necklace-shaped ;
cylindrical and with contractions at
intervals.
Monocarpellary. Of one carpel : 261.
Monocarpic (-icus). Monocarpous, Mon-
ocarpian. Only once fruiting; 33.
Monvcephalous (-us). Bearing a single
capitulum.
Monochasium. A cyme with one main
axis; 152, 155.
Monochlamydeous (-eus). Having but
one kind of perianth; 190, 340.
Monvoclinous (-us). Synonym of Herma-
phrodite.
Monocotyl don, Monocotylédones, adj.
Monocotyledonous (-eus). Plants or em-
brvo with a single cotyledon; 23, 27,
314, 339.

GLOSSARY.

Monolocular (-aris). One-celled.

Monecia. Name of Linnean class (335)
with flowers.

Monecious (-ius), or Monvicous (-us).
With stamens and pistils in separate
blossoms on the same plant; 191.

Monogamia, Name of a Linnzean arti-
ficial order, in class Syngenesia; 337.

Monograph. A systematic account of
parucular genus, order, or other group;
369.

Monogynia. Name of a Linnean arti-
ficial order, with solitary pistil, or
style; 337; hence, adjectively, Mond-
gynous ; 267.

Mondémerous. Formed of a single mem-
ber; 176.
Monopetdlous (-us). Literally one-

petalled; but always used in the
sense of Gamopetalous, which term is
to be preferred ; 244.

Monophyllous (-us). One-leaved.

Ménopode (Monopdédium), Monopodial.
A stem of a single and continuous
axis; 55.

Mondplerous (-us). One-winged.

Monepyrcnus. Containing a single stone
or nutlet. j

Monosepalous (-us). Equivalent to Gam-
osepalous; but literally of a single
sepal; 244.

Monospcrmous (-us).

Mi mnostichous (-us).
rank.

Monostylous (-us) With a single style.

Monosymmetrical. That which ean be
bisected into equal halves in only one
plane; 175.

Monétocous (-us). Bearing progeny
(fruiting) only once, as annuals and
biennials ; 33.

Monster (Monstrum). A monstrosity,
or unnatural development.

Morphology, 5.

Moschate (-atus). Exhaling the odor of
musk.

Mueilaginous, Mueilaginosus. Slimy;
of the consistence or appearance of
mucilage.

Mucro, Mucronation. A short and
abrupt small tip to a leaf, &e.

Micronate {-atus). Tipped with a mu-
cro; 97.

Mucrénulate (-atus).
nate.

Mule. A hybrid or cross-breed.

Multicipital ( Multiceps). Many-headed ;
many shoots or stems from the erown
of one root.

One-seeded.
In a single vertical

Minutely muero-

GLOSSARY.

Multifarious (-us). Many-ranked, as
leaves in several vertical ranks

Miultifid (-idus). Cleft into many lobes
or segments.

Multiflorous (-us).

Multijigate (Multijugus).
pairs or juga.

Multilocular (-aris).
several-celled.

Multiparous. Many-bearing; said of a
several-branched cyme; 152, 158.

Multiple fruits. The tructification of a
flower-cluster when coufluent into one
mass; 301.

Multiplication. Same as Augmentation ;
179, 200.

Multiserial (-alis), Multisertate (-atus).
In several series.

Miurvicate (-atus). Rough with short and
firm excresceices.

Muriculate (-atus). Minutely muricate.

Museariformis. Vly-brush-shaped.

Musciform (-ormis). Moss-like in ap-
pearaiice.

Muscoloyy. The botany of Mosses. But
is a hybrid word, and is replaced by

Many-flowered.
In many

Many-celled or

Bryology.

Miticous (-us). Pointless, blunt, awn-
less.

Myeclium. The filamentous vegetative

growth of a Fungus.
Mycology, Mycetology. The botany of
Fungi. q
Mycropyle. Micropyle misspelled.

Naked. Wanting some usual covering ;
as flowers without perianth, ovules
without coats, seeds not in a pericarp,
buds without seales.

Napiform (-ormis). Turnip-shaped ; 31.

Nanus. Dwarf.

Natant (-ans).
under water.

Navieular (-aris).
as Cymbiform.

Nebulose (-osus). Clouded or misty.

Neck. See Collum.

Necklace-shaped. See Moniliform.

Nectar. The sweetish secretion by va-
rious parts of the blossom from which
bees make honey.

Nectary (Necturium). The place or
thing in which nectar is secreted:
formerly applied also to any anoma-
lous part or appendage of a flower,
whether known to secrete honey or
not; especially to the hollow spurs of
a Violet, Larkspur. Columbine, and
the like.

Floating or swimming

Boat-shaped. Same

421

Nectariferous (-us). Nectar-bearing.
Needle-shaped. See Acerose.

Nemorosus, Nemoralis. Inhabiting
groves.
Nervation. Same as Venation, or un-

branched venation.

Nerve (Nereus). In botany, this is a
simple or unbranched vein, or a slen-
der rib.

Nerved, Nervose (-osus), Nervate (-atus).

Having ueryes in the botanical
sense.

Nervulose (-osus). Diminutive of ner-
vose.

Netled. Same as Reticulated; Netted-

veined; 92.

Neurose (-osus). Same as Nervose.
Neura being the Greek for nerve.
Neuter, Neutral. Sexless; as a flower
which has neither stamen nor pistil;

191, 195.

Niger. Black or blackish.

Nigricuns. Turning black or verging
to black.

Nitidus. Smooth and shining.

Nicalis. Growing in or near snow.

Niveus. Suow-white.

Nodding. Uanging down.

Node (Nodus). Literally a knot; the
portion of a stem which normally
bears a leaf or whorl of leaves; 6.

Nedose (-osus). Kuotty or knobby.

Nodulose (-osus). Diminutive of Nodose.

Nomenclature, 3, 345.

Normal (-alis). According to rule;
agreeing with type.

Notate (-atus). Marked by spots or lines.

Nothus. Yalse or bastard.

Notorhizal (-izus). Synonym of incum-
bent, as applied to the embryo of
Cruciferz.

Nuciform (-ormis). Nut-like in shape.

Nucleus. A kernel of an ovule, seed,
&e.; 277. A soft solid interior part
of a vegetable cell in the early condi-
tion; 309.

Nuculanium. Name given by Richard
to a drupaceous or baccate fruit con-
taining more than one stone or stony
seed; adopted by Lindley for a supe-
rior stony-seeded berry, such as a
grape.

Nucule (-ula). A diminutive nut or
stone; same as Nutlet; 296.

Nucum+ntaceus (-eus). Nut-like in char-
acter.

Nudicaulis. Naked-stamened ; stem not
leafy.

Nudus. Naked, in its various senses.

422

Numerous (Numerosus). Used in the
sense of indefinite in number.

Nut (Nuzx). A hard and indehiscent
one-seeded pericarp resulting from a
compound ovary; 295.

Nutlet. A diminutive Nut. See Nucule.

Nutant (-ans). See Nodding.

Ob. Over against; as a prefix denotes
inversely or oppositely, as

Obcompressed (-us). Flattened the other
way, antero-posteriorly instead of lat-
erally.

Obconical (-icus). Conical, but attached
at the apex.

Vbcordate (-atus). Inverted _heart-
shaped, the notch at the apex; 97.
Obdiplostemonous (-us), Obdiplostemony.
When the stamens are double the
number of the petals, but the outer

series opposite the latter; 198

Obimbricate. Imbricated or successively
overlapping downward.

Oblanceolute. Lanceolate but tapering
toward the base more than toward the
apex ; 95.

Oblique ( Obliquus).
slanting.

Oblong ( Oblongus). Considerably longer
than broad and with nearly parallel
sides; 95.

Obovate (-atus). Ovate with the broader
end toward the apex; 95.

Oboroid Solid obovate.

Obtectus. Covered by something.

Obtuse (-usus). Blunt or rounded at the
extremity ; 97.

Obtusiusculus. Somewhat obtuse ; dimin-
utive of obtuse.

Obvallatus. Guarded on all sides or sur-
rounded as if walled in.

Obverse, Obversely. Same as the prefixOb.

Obvolute (-utus). A modification of Con-
volute; 138, 139

Ocellate (-atus). With a circular patch
or a ring of color.

Ochraceous (-eus). Ochre color; light

_ yellow with a tinge of brown.

Ochrea, Ocrea. A leggin-shaped or
tubular stipule or rather combined
pair of stipules; 106.

Ochreate, Ocreate (-atus). Furnished
with ochrea or sheaths; 106.

Ochroleucous (-us). Yellowish-white or
between white and yellow.

Octo. Eight. In composition gives such
terms as the following.

Octacynia. Linnean artificial order with
Octagynous (eight-sty led) flowers ; 337.

Unequal-sided or

GLOSSARY.

Octamerous.
the circle.

Octandria. The Linnean class with
Octandrous, i. e. eight-stamened flow-
ers; 249, 334.

Octéni. In eights.

Octopetalous (-us). Eight-petalled. ~

Octosepalous. With eight sepals.

Octostichous (-us). In eight vertical
ranks; 124.

Oculate (-atus). Same as Occllate.

Officinal (-alis). Used in medicine or
the arts, therefore in the shops.

Offset. A short lateral shoot for propa-
gation; 53.

Oides, Oideus, -odes and -ides. Greek
for likeness, used as terminations, in-
dicate similarity to; as Dianthoides,
resembling a Pink.

Oleraccouz (-cus). Esculent in the way
_of a pot-herb.

Oligos. Greek for few; in compounds
giving such terms as

Oligandrous (-us). With few stamens.

Oliganthous (-us). Few-flowered.

Oliyémerous (-us). Of few members.

Oligospérmous (-us). Few-seeded.

Olivaceus (-eus). Olive-green.

Omphalodium. A mark (navel) on the
hilum of a seed, through which passed
vessels to the chalaza or rhaphe.

One-sided. Fither turned to one side,
or with parts all turned one way, or
unequal-sided. :

Oophovidium. The spore-case for the
larger spores in Selaginella, &c.

Opaque (Opdcus). Mostly used in the
sense of not shining or dull.

Operculate (-atus). Furnished with an
Operculum or lid.

Operculum. <A lid; a top which sepa-
rates by a transverse line of separation
as does that of a pyxis.

Opposite (-itus). Set against; as leaves
over against each other when there
are two on one node; or one part be-
fore another, as a stamen before a
petal; 6, 120, 178.

Oppositifolius (-ius). Placed opposite a
leaf, as is a tendril or peduncle in
Vitis, &e.

Oppositipetalous (-us).

Composed eight parts in

Placed before a-

petal.

Oppositisepalous (-us). Situated before
a sepal.

Orbicular (-arvis), Orbiculate (-atus).

Said of a flat body with a circular out-
line; 95.
Orchidacevus, 246.

GLOSSARY.

Order (Ordo). Group between genus (or
tribe) and class; 328.

Ordinal. Relating to orders.

Organdgeny (Organogenesis). The for-
mation or early development of or-
gans ; 2.

Organéography, Oryandlogy. The study
of organs and their relations; 2.

Organs of Vegetation, 11.

Orgyalis. Six feet high, or of the height
of a man. :

Ornithéphilous. Said of flowers which
are habitually fecundated by pollen
brought by birds; 217.

Orthoploceus. Said of an embryo when
incumbent cotyledons «re folded
around the radicle, as in Mustard

Orthos, Greek for straight; whence com-
pounds such as the following

Orthostichies. Vertical ranks; 121.
Orthostichous. Straight-rauked.

Orthétropous (-us), Orthotropal. De-
notes an ovule or seed with straight
axis, chalaza at the insertion, and ori-
fice at the other end; 277. Has been
applied to an embryo with radicle
pointing to the hilum; 312.

Os (oris). The mouth or orifice.

Osseous (-us). Of the texture of bone.

Ossiculus. A little stene, same as Pyrena.

Ostidlate (-atus). Furnished with a small
orifice or little door ( Ostiolum).

Outgrowths, 209.

Oval (Ovalis). Broadly elliptical; 95.

Ovary (Ovarium). The ovuliferous part
of a pistil; 166.

Ovate (Ovatus). Of the shape of the
longitudinal section of a hen’s egg,
the broader end basal; 95. Used also
for an egg-shaped solid.

Ovoid (Ovvideus). Used either for solid
ovate or solid oval, more properly for
the latter.

Ovulate (-atus), Ovuliferous.
ovules.

Ovule, Ovuluim. The body in the flower
which becomes a seed; 166, 276.

Bearing

Pagina. The surface of any flat body,
such as a leaf.
Palaceous (-eus). When the edges, as of
a leaf, are decurrent on the support.
Palate (Palatum). A projection in the
throat of a personate gamopetalous
corolla; 248.

Palea. A chaff, or chaff-like bract, such
as the chaffy scales on the receptacle
of the head in many Compositx; also

|

i

423

an inner bract or glume in Grasses;
142.

Paleaceous (-eus). Chafiy ; furnished
with pale; or chaff-like in texture.
Paleola. A diminutive palea, or one
of a secondary order; one of the
names of the Lodicule or Squamella
in Grasses.
Paleolate (-atus).

ole.

Palets (Pales of some Evglish botanists).
Same as Paleze; 142.

Palmaris. A palms breadth or length;
z. e. equalling the breadth of the four
fingers of the palm.

Palmate (-atus}. Lobed or divided
so that the sinuses point to or reach
the apex of the petiole or insertion;
101.

Palmately (Palmatim or Palmati-).
the palmate manner.

Pulmately veined, 93

Palmatifid (-idus), Palmatilobate, Pal-
matisect). Palmately cleft, lobed, or
divided.

Palminerved. Palmately nerved; 93.
Paludose (-osus), Palustiine (Palustris
or Paluster). Inhabiting marshes.
Pandurate (-atus),Panduriform (-ormis).

See Fiddle-shaped.

Panicle (Panicula). A loose compound
flower-c luster, such as is produced by
the branching of a raceme, or the ir-
regular branching of a corymb; 150.

Panicled, Paniculate (-atus). In a pani-
cled manner or borne in a panicle.

Pannosus, Panniformis. Having the ap-
pearance or texture of felt or woollen
cloth.

Papery, Papyraceous. Having the text-
ure of paper.

Papilionaceous (-eus). Butterfly-like;
applied to a peculiar polypetalous
corolla; 184. 246.

Papillar (-aris), Papillose (-osus), Papil-
late (-atus). Bearing or resembling
papille, minute nipple-shaped projec-
tions.

Pappiferous (-us),
Bearing a pappus.

Pappus. Thistle-down; thence applied
to various hairy tufts on akenes or
fruits; and thence to any production
or structure which takes the place of
the limb of the calyx on the akenes of
Composite; 192, 295

Papuliferous (-us), Papulose (-osus).
Covered w.th Papule, or small pim-
ples

Furnished with pale-

In

Puppose (-osus).

424 GLOSSARY.

Paracarpium. Unused name given to
an abortive pistil or carpel.

Puracorolla. A crown or internal ap-
pendage or deduplication of a corolla.

Parallel-nerved, veined, &e. Same as
Nerved; 91.

Parapetalous (-us). Said of stamers,
&c., which stand at each side of a
petal; 178, 201.

Pardphysis, pl. Paraphyses. Jointed
thread-like bodies, of no known func-
tion, accompanying the archegonia of
Mosses.

Parasitic (-icus). Growing on or in and
living upon another plant or even ani-
mal; 36.

Purastémon. Name rarely applied to an
abortive stamen or body in place of or
accessory to a stamen; same as Sta-
minodium.

Parastichies Secondary spirals in phyl-
lotaxy; 127.

Parenchyma. Common or soft cellular
tissue.

Parenchymatous. Of the nature of or
composed of parenchyma.

Paries, pl. parietes. The wall of any
organ.

Parictal (Parietalis). Borne on or re-
lated to the wall; 265.

Paripinnate (-atus). Even-pinnate ; same
as abruptly pinnate; 101.

Parted, Partite (-itus). Cleft nearly
but not guite to base; 98.

Parthenogénesis, Parthenogeny. Pro-
duction of seed without the interven-
tion of pollen; 285.

Partial (Partials). Secondary, as Par-
tial involucre (142), peduncle (143),
petiole (105), umbel (150), &c.

Partible (Partibilis). At length sepa-
rating or easily to be separated.

Partition. In one sense a separated por-
tion or segment; in another and the
more usual, a wall or dissepiment.

Patelliform (-ormis). Disk-shaped, cir-
cular with a rim, of the form of the
patella or kneepan.

Patent (Patens) Spreading ;_ either
widely open or diverging widely from
an axis.

Patentissimus. Superlative of Patens;
extremely spreading.

Patulous (-us) Slightly or moderately
spreading.

Pauciflorous (-us). Few-flowered.

Paucifolius. Few-leaved.

Petr-shaped. Obovoid or obeonical with
more tapering base.

Peéctinate (-atus). Pinnatifid with nar-
row and closely set segments, like
comb-teeth.

Pedilis A toot long or high.

Pedate (-atus). Palmately divided or
parted with the lateral divisions two-
cleft; resembling a bird's foot.

Pedatipartitus, -lobatus, -sectus, &e. Pe-
dately parted, lobed, divided, &e.

Pedicel (-clius). An ultimate flower-
stalk or its division; the support of a
single flower; 143.

Pedicellate (-atus). Pedicelled, borne
on a pedicel.

Pediculus. Name sometimes used for
Pedicel.

Pedunele (Pedinculus). A general
flower-stalk, supporting either a clus-
ter or a solitary flower; in the latter
case,the cluster may be regarded as
reduced to a single blossom; 143.

Pedinculate (-atus). Peduncled, borne
on a foot-stalk.

Pelovia. An irregular flower become
regular by a monstrous development
of complementary irregularities ; 186.

Peltate (-atus), LPeltiforvm (-ormis).
Shield-form ; target-shaped; a plane
body attached by its lower surface
(instead of margin or base) to a stalk;
96, 107.

Peltinerved (-ius) Radiately-nerved or
ribbed all round the circle.

Pelviform (-vrmis). Basin-shaped ; shal-
low cup-shaped.

Pendent (-cns). Hanging on its stalk or

support.

Pendulus (-us), Pendulinus. Hanging
more or less, as if from weakness of
the support.

Penicillate (-atus), Penicilliform (-ormis).
Pencil-shaped, the pencil (penicillum)
being a brush or tuft of hairs.

Pennute (-atus). Same as Pinnate.

Penniform (-ormis). In the form of a
feather or its plume.

Penninerved (-ervius). Same as pin-
nately nerved or veined; 93.

Penta. Greek for five; gives compounds
such as

Pentacarpellary. Composed of five car-
pels; 261.

Pentachenium. Name of a pentacarpel-
lary fruit otherwise like a cremocarp.
Pentadeélphous (-ws). With stamens in

five clusters; 250.

Pentacynia. Linnean. artificial order
characterized by Pentagynous, 7. €
five-styled flowers; 337.

GLOSSARY.

Pentamerous (-us). Composed of five
members in a circle; 176.

Pentandria. The Linnean class with
Pentandrous, i. e. tive-stamened flow-
ers; 249, 534.

Pentap:talous (-us). Five-petalled; 244.

Pentaphyllous \-us). Vive-leaved: 243.

Pentapterous (-us). Five-winged.

Pentasepalous(-us). Of five sepals; 244.

Pentastichous(-us). In tive vertical ranks ;
123.

Pepo, Peponida, Peponium.
fruit; 298. Z

Perennial (Perennis, Perennans). Last-
ing year after year; 32.

Perfect (Perfectus). Said of a flower
which is hermaphrodite.

Perfoliate (-atus). Where a stem seem-
ingly passes through a leaf; 167.

Perforate (-atus). Pierced, or having
translucent dots which look like holes.

Pergameneus, Pergamentaceus. Parch-
ment-like in texture.

Peri. Greek for around; hence such
compounds as

Perianth (Perianthium). The floral en-
velopes or leaves of the flower, consist-
ing of calyx, corolla, or both; 164, 243.

Pericarp (-arpium). The fructitied
evary; 286.

Pericarpic (-icus).
carp.

Perichetial (-ialis). Relating to the
Perichetium, a set of bracts around
the fruit-stalk in Mosses.

Pericladium. The sheathing base of a
leaf when it expands and surrounds
the supporting branch.

Periclinium. Involucre of the capitu-
lum of Composite; 148.

Péviderm (-ermu or -ermis)
or Epiphlceum.

A gourd-

Relating to the peri-

Outer bark

Pévigone, Perigénium. Synonym of
Perianth; 164.
Perigynium. Name of hypogynous

bristles, scales, or a sac, which sur-
rounds the pistil (also the stamens
when present) of many Cyperacez.
Perigynous (-us} Literally around the
ovary: said of organs which are ad-
nate to the perianth, or to this as con-
nate with the lower part of the pistil;
182.
Peripétalous (-us). Around the petals.
Peripheric (-icus). Of or belonging to the
circumference ; as of an embryo coiled
round the outside of the albuinen.
Periphoranthium. Synonym of the
involucre of Composite; 148.

425

Peripterus. Surrounded by a wing or
thin border.

Perisperm (-ermium). The albumen of
the seed, at least the exterior and or-
dinary albumen; 14, 310.

Peristome (Peristoma or Peristémium).
The fringe or other structure surround-
ing the orifice (stoma) of a Moss.

Peritropous (-us) or Pevitrepal. Said
of a seed which is horizontal in the
pericarp; or of a radicle pointed to-
ward the sides of the pericarp.

Persistent (-ens). Remaining even on
the fruit, or over winter; 243.

Peérsonate (-atus) Masked, as when a
bilabiate corolia has a prominent pal-
ate; 248.

Pervious (-ius).
way.

Pertuse (-usus). Having slits or holes.

Perula, pl. Perule. Scales of leaf-buds
and the like; 40.

Perulate (-atus). Furnished with peru-
le or scales.

Pes, gen. pedis. A foot. Hence in
Latin compounds Lonyg/pes, long-
stalked, Brevipes, short-sialked, &e.

Petal (Petalum). <A corolla-leaf; 165.

Petaline (-inus), Petalvid (-oideus).
Petal-like, or relating to petals; 118.

Petalody. Name for the metamorphosis
of other organs (such as stamens) into
petals; 174.

Petiolar (-aris).
to a petiole.

Petiolate (-atus), Petioled.
petiole.

Peétivle (Petiolus).
leaf; 85, 104.
Petidlulate (-atus),

Having a

Pétiolule (Petidlulus).
leaflet; 105.

Petreus. Growing among rocks.

Petrosus. Growing in stony places.

Phenogams, Phenogamia, Phenoga-
mous plants. Plants sexually propa-
gating by flowers, of which the essen-
tial organs are stamens and pistil; 3,
334, 340, 344.

Phalanges, sing. Phalanz. The bundles
of stamens in diadelphous or polyadel-
phous flowers.

Phanerogams, Phanerogamia, &e. See
Phenogams, «ce.

With an open passage-

Borne on or relating
Having a
The footstalk of a
Petiolular

(-a7ris).

A footstalk of a

Phleum. Greek name for bark.

Pheniceus. Deep red with some sear-
let.

Phoranthium. A name for the recep-

426

tacle of the capitulum in Composite ;
148.

Phycology. The botany of Algze.

Phylla. Leaves in Greek; combined
with Greek numerals, forming such
terms as Diphyllous, Triphyllous, &c.,
to Polyphyllous. :

Phyllocladium. A branch assuming the
function of foliage; 65.

Phyllodineous (-eus). Relating to a

Phyliidium. A petiole usurping the
form and function of a leaf-blade;
110.

Phyllody, Phyllomorphy. Names for
the transformation or metamorphosis
of floral organs into leaves; 174.

Phyllotaxis, Phyllotary. Leaf-arrange-
ment; 119.

Phyllomania. The unusual or abnormal
production of leaves.

Phyllophore (-orum). The budding sum-
mit of a stem on which leaves are de-
veloping.

Phyllum.
Phylla.

Phyllome, Phylloma. An assemblage
of leaves, or of incipient leaves in a
bud. Also recentiy used by German
botanists for leaf generically or poten-
tially, that which answers to a leaf; 6.

Phytography. Botany as relates to the de-
scription and illustration of plants ; 345.

Phy dlogy. Synonym of Botany.

Phiytomer, pl. Phytomera. Plant-ele-
ments in morphology ; same as

Phyton. Greek name for plant; has
been used in the sense of plant-ele-
ment, or plant-unit; 7.

Phytotomy. Same as Vegetable Anatomy
or Histology; 2.

Piceus. Pitch-black or brownish-black.

Pictus. Painted, or rather as if painted.

Pileate (-atus), Pileiformis. Having the
form of a cap or Pileus.

Pileorhiza. The root-cap.

Piliferous (-us). Bearing or tipped with
hairs (pili).

Pilusciusculus. Slightly hairy.

Pilose (-osus). Hairy, in general with
any sort of pilosity ; in particular with
soft and distinct hairs.

Pinna. One of the primary divisions of
a pinnate leaf, either simply pinnate,
when it is a leaflet, or a partial petiole
or rhachis with the leaflets when the
leaf is bipinnate; 104

Pinnate (-atus). When leaflets are
arranged along each side of a com-
mon petiole ; 100.

Greek for leaf; 6, 85. See

GLOSSARY.

Pinnately cleft, lobed, parted, &e.; 99.

Pinnately veined. Yeather-veined; 93.

Pinnatifid (-idus). Pinnately cleft. -

Pinnatilobatus, Pinnaidobus. Pinnately
lobed.

Pinnatipartitus. Pinnately parted.

Pinnatisictus. Pinnately divided quite
down to the rhachis.

Pinnule (Pinnula). One of the pinnately
disposed divisions of a pinna; a sec-
ondary pinna; 104.

Pisiform (-ormis). Pea-shaped; resem-
bling a pea.

Pistil (Pistiilum). The female organ‘ of
a flower, consisting of ovary, style.
aud stigma, or at least of ovary and
stigma; 302, 259.

Pistillate (-atus), Pistilliferous. Said of
a plant or a blossom provided with
pistil, most properly for one having
pistilonly; 191.

Pistillidium. One of the names of the
analogue of pistil in Mosses, &e.

Pistillody. Name for the metamorpkosis
of other organs into carpels; 174.

Pitcher. See Ascidium. A tubular or
cup-shaped leaf, which usually holds
some liquid; 111.

Pith. A ceutral cellular part of a stem,
especially of an exogenous stem; 75.

Pitted. Marked with small depressions
or pits.
Placenta. That in the ovary which

bears the ovules, sometimes the mere
united margins of the carpel-leaves,
somctimes a thickening or enlarge-
ment of them, or even of some other
part of the ovary; 261.

Placentation (-iv). The disposition of
the placentz.

Placentiform (-ormis). Quoit-shaped,
or in form like a flat cake.

Plaited. See Plicate.

Plane (Planus). With flat surface or
surfaces.

Platys. Greek for wide, in such com-
pounds as Platyphyllus, broad-leaved,
&e. ;

Pleios. Greek for full, used in com-
pounds for several or many; as Pleto-
phyllous, several-leaved, &e. Simi-
larly Pleistos for a great many.

Pleiockdsium. A several-rayed cyme;
152, 155.

Plenus. Full. Flos plenus is what gar-
deners call a ‘‘ double flower,’ that is
one in which the petals or other flower-
leaves are abnormally multiplied.

Pleur-nchymat. Same as woody tissue.

GLOSSARY. 427

Pleurorhizal (-us). Embryo with radicle
against one edge of the cotyledons;
z. e. the latter accumbent.

Plicate (-atus), Plicativus. Folded into
plaits (plice), usually lengthwise; 133,
139.

Plimbeus. Lead-colored;
with some metallic lustre.

Plumose (-osus). Feathered; when bris-
tles, &ec., have fine hairs on each side
like the plume of a feather, as the pap-
pus of Thistles.

Plumule (Plumula).
ing point of the embryo above the
cotyledons; 17.

Plures. Many or several; used as a
prefix in Latin words, such as Pluri-
florous (-us), several-tlowered ; Pluri-
loculur (-aris), several-celled ; Pluri-
Joliolate, with several leaflets, &c.,
Plurijugate, in several pairs, &e.

Péculiform (-ornis). In the shape of a
drinking-cup or goblet.

Pod. A dry and several-seeded dehis-
cent fruit; strictly a Leguine or a
Silique ; 288, 292.

Podium, Podus. A footstalk, stipe, or
other such support; used only in Greek
compounds, as Podocephalus, head
pedunculate; Podocarpus, fruit stipi-
tate; or as a suffix, in such words as
Leptopodus, slender-stalked; Brachy-
podus, short-stalked, &c.

Podetium. Any stalk-like elevation.

Podogynium. Same as Gynophore.

Podosperm (-ermium). The stalk of a
seed; 276, 305.

Pogon. Greek fora beard ; enters into
various compound words.

Polémbryony. See Polyémbryory.

Politus. Polished; applied to a smooth
and shining surface.

Pointless. Same as Muticous.
Pointletted. Minutely pointed; same as
apiculate or as minutely acuminate.
Pollen, Pollen-grains. The fecandating
grains or cells contained in the anther;

165, 256.

Pollen-tube. The slender tube which
begins as a protrusion of the inver
coat of a pollen-grain, and elongates
by growth, at least when in contact
with the stigma; 258.

Pollicaris. An inch long; the length
of the terminal joint of the thumb,
pollex.

Poliniferous (-us). Pollen-bearing.

Pollinium. A mass of pollen-grains
more or less coherent ; 257, 230.

dull gray

The bud or grow-

|

Pollinated (-1tus)}. Said of a stigma
when supplied with pollen.

Poly. In Greek compounds, denotes
numerous ; as in

Polyadelphia. Name of a Linnzean ar-
titicial order with stamens Polyadel-

phous, or in several phalanges or
brotherhoods; 250, 335.
Polyandria. Name of a Linnean class

with flowers Polyandrous, or having,
an indefinite number of stamens; 249.
334.

Polydnthous (-us). Many-flowered;. in
the Latin form same as multiflorous.
Polycarpellary. Of many carpels; 261.
Polycarpic (-icus). Fruiting many times
or indefinitely; DeCandolle’s name

for a perennial herb; 33.

Polycephalous (-alus). Consisting of or
bearing many heads, eupitula.

Polycoccus. Of several cocci.

Polycotyledonous (-eus oF es).
several cotyledons; 22, 314.

Polygamia. Name of a Linnzean class
having Polggaimous flowers, i. e. some
hermaphrodite, some unisexual ; 191,
335. Also of Linnean orders of Syn-
genesia; 337.

Polygynia. Name of a Linnzean artifi-
cial order with flowers Polygynous,
7. e. containing numerous carpels;
261, 337.

Polymerous (-us). Of numerous mem-
bers to each series or circle.

Polymorphous (-us). Of several or vari-
ous forms.

Polypetalous (-us).
petals; 244. :

Polyphore (-drium). A torus which
bears many pistils, as that of a straw-
berry or raspberry} y-

Polyphyllous (- -us). Many-leaved; 244.

DP lysepalous (-us). Of separate sepals;
244

Polyspérmous (-us).

Polystemonous (-us).
mens.

Polystachyus. Bearing many spikes.

Polystylous (-us). Bearing many styles.

Polysymmetrical. That which can be
divided into similar halves in several
or more than one plane; 175.

Polytocous (-us). Bearing progeny (fruit-
ing) many times, 7. c. year after year;

Pome (Pomum). Kind of fruit of which
the apple is the type; 298.

Pomeridianus. In the afternoon.

Pomiferous (-us)., Pome-bearing. __.

Having

Having separate

Many-seeded.
With many sta-

428

Pomology. A treatise on or the subject
of fruits considered as esculent.

Porouse (-osus), Porous. Pierced with
small holes or pores.

Posterior. In an axillary flower is the
side next the axis of inflorescence;
160.

Posticous (-us). On the posterior side,
which in a flower is that next the axis
of inflorescence: an adnate anther is
posticous when on the outer side of
the filament, 7. e. when it faces the
petals; 253.

Pouch. See Silicle.

Precot. Appearing or developing early

Prefloration. Same as Hstivation; 132.

Prefoliation. Same as Vernation; 132.

Premorse (-orsus). With end as it were
bitten off.

Prasinus. Grass green.

Pratensis. Growing in meadows.

Prickly. Armed with Prickles (56),
which are outgrowths of the bark or
rind.

Primine. Outer coat of the ovule; 277.

Primordial (-ialis). The first in order
of appearance. Primordial leaves are
those of the plumule.

Prismatic (-icus). Prism-shaped, with
flat faces separated by angles.

Procerus. Very tall.

Process (Proccssus). Any projecting
appendage.

Procumbent (-ens).
ground; 53.

Productus. Produced, 7. e. extended or
prolonged into.

Pro-embryo, 284.

Proles. Progeny; sometimes used for
race; 32i.

Proiiferous (Prélifer, Proliferus). Bear-
ing progeny, in the way of offshoots.

Proliferation or

Prolification is usually taken as the
production by one organ of something
different, such as the development of
buds and plantlets on leaves, of leafy
shoots in place of flowers, &c.: 73.

Proligerous (-us). Same as Proliferous.

Prone (Pronus). Lying flat, especially
face downward.

Propaculum, Propagulum. Name of a
shoot, such as a runner or sucker which
may serve for propagation.

Propdgines. Same as Bulblets.

Prophylla. Primary leaves, as the first
leaves of a b-anch or axis.

Prosenchyma. Plant-tissue consisting of
lengthened, tubular, or fasiform cells.

Lying along the

GLOSSARY. .

Prostrate (-atus).
the ground; 53.

Protos. Greek for first; used in various
compounds, such as

Protundrous, Protand:y.
androus.

Proterdndrous, also Protandrous, Pro-
terandry. When the anthers of a
flower are in alithesis earlier than the
stigma; 219, 220.

Proteranthous (-us).
precedes leafing.

Proterogynous, Proterogyny, or Pro-
togynous, Protogyny. When the
stigma is ready for its functious ear-
lier than the anthers of the same
blossom; 219.

Protophytes, Pretophyta.
the supposed first plants.

Protoplasm, Protoplasma. The forma-
tive organic material of plauts and
animals, in its living state.

Pruinate (-atus), Pruinose (-osus). As if
frosted over with a bloom or powder.

Pseudos. Greek for false, a prefix in
various compounds, as Pseudo-mono-
cotyledonous ; 26.

Pseudo-bulb. A thickened and bulb-
like internode in epiphytal orchids; a
corm.

Pseudocarp (-arpium). The principal
or accessory part of an anthocarpous
fruit; 300.

Pseudo-costate False-ribbed, as where
a marginal or intramarginal vein or
rib is formed by the confluence of thie
true veins.

Pseudospermium. Name given to any
kind of «ne-seeded fruit which is inde-
hiscent and resembles a seed, such as
an akene, &e

Psilos. Greek for naked or bare; as in
Psilostachyus, with naked spike.

Pteridium, Pteiodium. Names for the
Key-fruit or Samara.

Pteris. Used for wing in Greek com-
pounds, also for a Fern.

Pteridographia. The botany of Ferns.

Pterocarpous (-us). Wing-fruited.

Pteropodus. Wing-footed, t. e. petiole
wing-inargined, &c.

Ptyzis. Greek name for folding, as of
leaves in a bud; 132, 133.

Pubens, Pubes Used for Pubescent.

Pub*rulus. Minutely pubescent.

Pubes. Pubescence, hairiness.

Pubescent (-ens). Clothed or furnished
with hairs or down, especially with soft
or downy and short hairs.

Lying quite flat cn

See Proter-

Where flowering

Alga, &c.,

GLOSSARY.

Pugioniform (-ormis). Dagger-shaped.

Pullus. Dark-colored; dusky-brown or
blackish.

Pulvereus, Pulverulentus. Powdered; as
if dusted with powdery matter or
minute grains.

Pilvinate (-atus), Pulviniform (-ormis).
Cushion-shaped.

Pulvinus. A cushion; name given to an
enlargement or swelling close under
the insertion of a leaf, or sometimes
to the swollen base of a petiole.

Pimilus. Low or little.

Punetate (-atus). Dotted, either with
depressions like punctures, or trans-
lucent internal glands, or with colored
(lots.

Puneticulate (-atus). Minutely punctate.

Pungent (-ens). Terminating in a rigid
and sharp point or acumination, like a
prickle.

Puniceous (-eus). Bright carmine-red.

Purpureus. Originally the red of arte-
rial blood; but our purple is some-
what dull red with a dash of blue or
violet.

Purpurascens. Purplish.

Pusillus. Very small, or weak and slen-
der.

Pustular, Pustulate (-atus),
(-osus).
blisters.

Putdmen. The shell of a nut; the endo-
carp of a stone-fruit; 288.

Pyenos, Greek for thick; whence Pyenv-
cephalus, thick-headed, &c.

Pygmeus. Dwarf, pygmy.

Pyramidal (-alis). Pyramid-shaped.

Pyréne (Pyréna). Same as Nucule or
Nutlet; one of the small stones of a
drupaceous fruit: 298.

Pyrenarium, Pyridium. A pear or pear-
like fruit, same as Pomum.

Pyrenarvius. Name of a drupaceous
pome, as of Medlar and Crategus.

Pyridion. Synonym of Pome.

Pyrenocarp (-arpium). A general name
for any drupaceous fruit; 292.

Pyriform (-ormis). See Pear-shaped.

Pyzxidate (-atus). YVurnished with a lid.

Pyzidium, Pyxis. A capsule with trans-
verse dehiscence, making a lid of the
upper portion; 293.

Pustulose
Having low elevations, like

Quadri-. In Latin compounds, denotes
four; as Quadrangular, Quadrifiri-
ous (in four vertical ranks), Qudriju-
gate (in four pairs), &c.

429

Quaternary, Quaternate. In fours or
composed of four; 176.

Quini, Quinary (-ius), Quinate (-atus).
In fives; 176.

Quinqgue. Five. In Latin compounds,
giving rise to such terms as

Quincuncial, in a Quincunz; also five-
ranked; 125, 156.

Quinquefarious (-ius),
ranks.

Quinquefoliate (-atus). Five-leaved.

Quinquefoliolate, with five leaflets.

Quintuple. Dividing into five parts, or
five-fold.

Quintuplinerved or -veined. With mid-
rib of leaf dividing into five (i.e. two
lateral pairs) above the base; 93.

In five vertical

Race. A variety of such fixity that it
is reproduced by seed; also used ina
looser and more extended sense for a
series of related individuals without
particular regard to rank; 320.

Raceme (Racemus). An indeterminate
or centripetal form of inflorescence
with lengthened axis and equal-pedi-
celled flowers; 146.

| Racemiferous. Bearing racemes.

Racemiform (-ormis). In the form of a
raceme.

Racemose (-osus). Having the character
or appearance of a raceme, or in ra-

ceines.

Rachis. See Rhachis
Radial. Belonging to the ray.

Radiate (-atus). Spreading from or
arranged around a common centre,
or around the circumference of a cir-
cle; bearing rays or ray-flowers.

Radiately veined. Same as Palmately
veined; 93.

Radiatiform (-ormis). Said of a eapitu-
lum of flowers which is radiate by en-
largement of some of the outer flowers,
which however are not truly ligulate,
as in species of Centaurea.

Radical (-alis). Belonging to or pro-
ceeding from the root, or from a root-
like portion of stem at or below the
surface of the soil.

Radicant (Radicans). Rooting.

Radicel. A minute root or a rootlet.

Radiciflorous (-us). Flowering (appar-
ently) from the root.

Radiciform (-ormis), Radicinus. Of the
nature or appearance of a root.

Radicle (4eula). Literally a diminutive
root; but the “‘radicle’’ of the embryo,

450

so called in descriptive botany, is the
hypocotyledonary and primal inter-
node. (See Caulicle); 10.

Radiculose (-osus). Bearing rootlets.

Rudiz. The root.

Rameal (-alis), Rameus.
(Ramus) a branch.

Ramenta. Thin chaffy scales belonging
to the surface or epidermis, such as
the chaff on the stalks of many Ferns.

Ramification. Branching; 47.

Ramijlorous (-us). Flowering on the
branches.

Ramose (-osus). Branching or branchy.

Ramulose (-vsus). Bearing many branch-
lets, ¢. e. Ramuli or

Riphe. See Rhaphe.

Rephides or Rhaphides. Crystals in
ihe cells of plants, especially needle-
shaped crystals.

Ray (Radius). One of the radiating
branches of an umbel (147); also the
niarginal as opposed to the central part
(or disk) of a head, umbel, or other
flower-cluster, when there is a differ-
ence of structure. Also used as an
abbreviated expression for

Ray-flowers. Those which belong to
the margin of a circular flower-clus-
ter, and differ from (being usually
larger than) those of the disk.

Recaulescence. The adhesion of leaves
or their stalks to a stem; 158.

Receptacle (Recepiaculum). A portion
of axis forming a common support or

bed on which a cluster of organs is
borne. The receptacle of the flower,
or the torus, is the axile portion of a
blossom, that which bears sepals,
petals, stamens, and pistils; 167, 211.
The receptacle of inflorescence is the
axis or rhachis of the head, spike. or
other dense cluster; 143.

Reclinute (-atus\, Reclined, Reclining.
Falling or turned toward downward,
so that its upper part rests on the
ground or other object; 53, 133.

Rectinervius. Straight-veined or straight-
nerved; 92.

Rectiserial (-ialis). In rectilinear ranks ;
124.

Recurved (-us), Recurvatus.
backward or downward.

Reduplicate (-atus) or Reduplicativus.
Folded and projecting outward.

Reflexed (-us). Abruptly bent or turned
downward or backward.

R. fracted (-us). Same as reflexed, but
abruptly bent from the base.

Belonging to

Curved

GLOSSARY.

Reygm1. A two-several-lobed two-
several-celled fruit (2-pluricoccous),
which separates at maturity into as
many 2-valved carpels, as in Euphor-
bia; one form of Schizocarp.

Regmacarp (-arpium). A general name
of a dry and dehiscent fruit, 292.

Regular (-aris). Uniform in shape or
structure; symmetrical as respects
shape; 175.

Reniform (-ormis). Kidney-shaped;
having the outline of the longitudinai
section of a kidney; 96.

Repand ( Repandus). With slightly un-
even margin, which, if more pro-
nounced, would be sinuate; 98.

Repent (Lepens). Creeping, 7. €. pros-
trate or horizonal and rooting; 53.

Replicate (-atus), Replicativus. Yolded
backwar.!.

Replum. <A frame-like placenta (like
a door-case), from which the valves
of a capsule or other dehiscent fruit
fall away in dehiscence, as in Cruci-
fere, certain Papaveraceez, Mimosa,
&c.; 293.

Reptant (Reptans). Same as Repent.

Resupinate. Upside down, or having
that appearance.
Rete. Network.

Reticulated (-atus), Retiformis.
form of network; netted.

Reticulate-veined, 92.

Retindculum. Name sometimes applied
to the gland to which one or more
pollinia are attached in Orchids, &e.
The persistent and indurated hook-
like funiculus of the seeds in most
Acanthacee.

Retinerved (-ius).
veined.

Retrocurved (-us). Same as Recurved.

Retroflexed (-us). Same as Reflexed.

Retrorse (-orsus). Directed backward or
downward.

Retroverted (Retroversus). Inverted.

Retuse (Retusus). With a shallow or
obscure notch at a rounded apex; 97.

Reversion A changing back, or in the
reverse direction: 171.

Rerolute (-utus). Rolled backward from
the margins or apex; 133.

Rhachis. The axis (backbone) of a
spike or of a compound leaf; 101, 143.

Rhaphe. The adnate cord or ridge
which in an anatropous ovule con-
nects the hilum with the chalaza;
279, 307.

Rhipidium. A fan-shaped cyme; 156.

In the

Same as Reticulate-

GLOSSARY. 43

Rhizanthous (-us). Root-flowered ; flower-
ing from the root or seeming root.

Lhizina. The peculiar roots or root-
hairs of Mosses, Lichenes, &c.

Rhizocarpous (-us). Rhizocarpic (-icus).
Literally root-fruited; used by De-
Candolle for a perennial herb.

Rhizome, Rhizima. A _ rootstock; a
stem of root-like appearance pros-
trate ou or underground, from which
rootlets are seut off; the apex pro-
gressively sending up herbaceous
stems or flowering stalks and often
leaves; 56. 7

Rhizomorphous (-us). Root-like in ap-

pearance.
Rhombic (-icus). Rhomb-shaped.
Rhomboidal (-alis). Approaching a

rhombic outline; quadrangular, with
the lateral angles obtuse.

Rib. A primary and strong vein or
conspicuous portion of the framework
of a leaf; 92.

Ribbed. Furnished with prominent
ribs.
Rictus. The mouth or gorge of a bila-

biate corolla.

Rima. A chink or cleft.

Rimuse (-osus). With chinks or cracks,
like those of old bark.

Ring. In Ferns, &c. See Annulus.

Ringent (-ens). Grinning or gaping ; as
is the mouth of an open bilabiate
corolla; 248.

Riparius. Growing along the banks of
rivers, &c.

Rirdlis Growing along brooks.

Rirvularis. Growing in watercourses or
rivulets.

Root. The descending axis. Roots are
axes which grow in the opposite di-
rection from the stem, are not com-
posed of nodes and internodes, are
mostly developed underground, and
absorb moisture, &c., from the soil;
2G

Root-cap, 13, 28.

Root-hairs. Attenuated unicellular
outgrowths or hairs from the newly
formed parts of a root, for absorp-
tion; 13, 29.

Rootlet. A very slender root or branch
of a root.

Rootstock. See Rhizoma; 56.

Roridus. Dewy; covered with particles
resembling drops of dew.

Rosaceous (-eus). Arranged like the
five petals of a normal rosé; 246.
Sometimes used for rose-color.

Roseus. Rose-colored; pale red.

Rostellate (-atus). Diminutive of Ros-
trate.

Rostéllum. A diminutive beak. Also

the name applied by Linneus to the
Caulicle or Radicle.

Rostrate (-itus). With a Rostrum, a
beak or spur; narrowed into a slender
tip or process.

Rosular, Rosulate (-atus).
a rosette.

Rotate (-atus). Wheel-shaped; circular
and horizontally spreading very flat ;
248.

Rotund (Rotundus, Rotundatus). Round-
ed in outline: 95.

Rough, Roughish, See Scabrous.

Rubellus, Rubescent (-ens), Rubens. Red-

Collected in

dish. Rubescent also is turning red.
Ruber. Red in general.
Rubicundus Blushing, turning rosy-
red.

Rubiginose (osus). Brownish rusty-red.

Ruderal (-alis). Growing in waste places
or among rubbish.

Rudiment. An imperfectly developed
and functionally useless organ; a
Vestige.

Rufous (-us), Rufescent (-ens).
red mixed with brown.

Rugose (-osus). Covered or thrown into
wrinkles, Ruge.

Ruminated (-atus). As if chewed; said
of the albumen of a nutmeg, &c.; 311.

Runcinate (-atus). Saw-toothed, or
sharply incised, the teeth or incisions
retrorse.

Runner. A prostrate filiform branch
which is disposed to root at the end
or elsewhere ; 53.

Running. Same as Repent.

Rupestris, Rupicola. Growing on rocks
or in rocky places.

Pale

Ruptilis. Bursting irregularly.
Rusty. Same as Rubiginose, Rufescent,

and Ferruginous.

Rutiluns. Deep red with a metallic
lustre.
Sabulosus. Growing in sandy places.

Saccate (-atus), Sacciform. Sac-shaped ;
baggy.

Sagittate (-atus), Sagittiform (-ormis).
Arrow-head-shaped.

Salsuginosus. Growing within reach of
salt water.
Salver-shaped.

phous; 248.

See Hypocraterimor-

452

Samara.
294.

Samaroid. Resembling a samara.

Sap-wood. New wood of an exogenous
stem; 80.

Sarcocarp (-arpium). The succulent or
fleshy portion of a drupe; 285. Has

een proposed also as a general name
for a baccate fruit; 292.

Sarmentose’ (-osus). Producing long
and lithe branches or runners, viz.
Sarments (Sarmenta).

Sativus. That which is sown or planted.

Saw-toothed. See Serrate.

Saxdatilis, Suxosus, Saxicolus. Living on
or among rocks.

Scabridus, Scabriusculus.
diminutive of

Seabrous (Scaber). Rough to the touch.

Scalariform (-ormis). Ladder-shaped;
with transverse markings like the
rounds of a ladder.

Scales. Any thin scarious bodies, usu-
ally degenerate leaves, sometinves of
epidermal origin.

Scalloped. Same as Crenate; 98.

Scaly. See Scarious, Squamoese.

Scaly Buds, 40.

Scandent (-ens).
ever mode; 51.

Scape (Scapus). A peduncle rising from
the ground; 51, 143.

Scapiform (-ormis), Scapose (-osus). Re-
sembling a scape.

Scapigerous (-us). Scape-bearing.

Scar. The mark left on the stem by
the separation of a leaf, or on a seed,
&c.; by its detachment.

Scarious or Scariose (-osus). Thin, dry,
membranaceous, and not green.

Schizocarp (-arpium). <A pericarp which
splits into one-seeded pieces; 296.

Scion. A young shoot; a twig used for
grafting.

Sciuroideus. Like a squirrel’s tail.

Scleranthium. Name of the fruit of
Mirabilis, and the like; an akene
enclosed in an indurated portion of
calyx-tube.

Scleroideus. Having a hard texture;
from Scleros, hard.

Scobiform (-ormis). Having the appear-
ance of sawdust.

Seorpioid. A form of unilateral inflo-
rescence which is cireinately coiled in
the bud: in the stricter sense, a form
with the flowers two-ranked, these
being thrown alternately to the right
and left; 155, 157.

An indehiscént winged fruit;

Roughish;

Climbing, in what-

GLOSSARY.

Scrobiculate ( atus). Marked by minute
or shallow depressions.

Scrotifurm (-ormis). Pouch-shaped.

Scurf. Small and bran-like scales on
the epidermis.

Scutate (-atus), ~Scutiform (-ormis).
Buckler-shaped.

Scutelliform (-ormis). Platter-shaped.

Scymetar-shaped. See Acinaciform.

Sectile (-ilis). As if cut up into portions.

Section (Sectio). In classification, is
applied in a general way to a divi-
sion in the arrangement of genera,
species, or other groups; 327.

Sectus. Completely divided; 99.
Secund (Secundus). When parts or
organs are all directed to one side.
Secundiflorus. With flowers of a cluster
all secund. :
Secundine. The second (inner) coat

of an ovule; 277.

Seed. The fertilized and matured ovule;
the result of sexual reproduction in a
phenogamous plait; 305.

Seed-leaves. Cotyledons, 11.

Seed-stalk. See Funiculus and Podo-
sperm.

Seed-vessel. See Pericarp.

Seyetalis. Growing in grain-fields.

Segment (Segmentum). One of the

divisions into which a plane organ,
such as a leaf, may be cleft.

Segregate (-atus). Separated; kept
apart.
Semen. Seed.

Semi. Half, in Latin compounds: such as

Semi-adherent. The lower half adhe-
rent, &e.; Semi-amplezicaul (-aulis),
half clasping the stem ; Semiovate,
ovate halved lengthwise, &c.

Semiandtropous. Same as Amphitro-
pous: 279.

Semilunar, Semilunate (-atus). A syn-
onym of Lunate, being like a half-
moon.

Seminal (-alis).

Seminiferous (-us).

Sempervirent (Sempervirens).
green.

Senary (-arius).

Sepal ( Sepalum).

Sepaline (-inus), Scpalous.
sepals.

Sepaloid (-oideus). Resembling a sepal.

Sepalody. Name for the metamorphosis
of petals, &e., into sepals or sepaloid
organs; 174. :

Separated flowers. Those of distinct
sexes; same as Diclinous; 191.

Relating to the seed.
Seed-bearing. -
Ever-

In sixes; 176.
A calyx-leaf; 165.
Relating to

GLOSSARY.

Septate (-atus).
tion or septum.

Septicide, Septiculal (-cidus). When a
capsule dehisces through the dissepi-
ments or lines of junction ; 289.

Septiferous (-us). Bearing the partition
or dissepiment.

Septifragal (-us). Where the valves in
dehiscence break away from the dis-
sepiments ; 290.

Septum. Any kind of partition, whether
a proper dissepiment or not,

Septulate (-atus). Divided by spurious
or transverse septa.

Serial (Serialis) or Seriate (Seriatus).
Disposed in series or rows, whether
transverse or longitudinal.

Sericeous (-cus). Silky; clothed with
close-pressed soft and straight pubes-
cence.

Serotinous (-us). Produced compara-
tively late in the season.

Serr:te (-atus). Beset with antrorse
teeth; 97.

Serrulate (-atus). Serrate with
small or fine teeth; 97.

Sesqui. <A Latin prefix denoting one
and a half; as, Sesquipedalis, a foot
and a half.

Sessile (-ilis). Sitting close, without a
stalk ; destitute of peduncle, pedicel,
or petiole, as the case may be.

Sets. A bristle, or bristle-shaped body.

Sctaceous (-eus). Bristle-like.

Setiform (-ormis). In the form of a
bristle.

Setigerous (-us). Bristle-bearing.

Setose (-osus). Beset with or abounding
in bristles ; bristly.

Setula. Diminutive of Seta.

Setulose (-osus). Bearing or consisting
of minute bristles.

Sex. Latin for six; as in Sexangular,
Sexfarious, Sexpartile, &c.

Shaggy. Pubescent with long and soft
hairs; same as Villous.

Sheath. A tubular or enrolled part or
organ, such as the lower portion of
the leaf in Grasses. See Vagina.

Sheathing. Enclosing as by a sheath.

Shield-shaped. In the form of a buckler;
plane and round or oval, with stalk
attached to some part of the under
surface; 96. See Clypeate, Scutate,
Peltate.

Shrub. A woody perennial of less size
than a tree: 50.

Shrubby. Having the character of a shrub.

S/eve-cells, 77.

Separated by a parti-

very

435

Sigillate (-alus). As if marked with the
impression of a seal, as the rootstock
of Polygonatum.

Sigmoid (-oideus). Doubly curved like
the Greek s or the capital S.

Silicle (Silicula). A short silique, not
very much longer than wide; 294.

Sil culusa. Name of the Linnean arti-
ficial order of the class Tetradynannia,

having Siliculose pods ; 337.
Silique (Siliqua). The peculiar pod of
Crucifere, especially when mech

longer than wide; 293.

Siliquosa. Name of the other order of
Tetradynamia, with Siliquose fruit,
i.e. a Silique; 337.

Silky. See Sericeous

Silver-grain. The glittering plates in
exogenous wood belonging to the
medullary rays; 74.

Simple (Simplex). Of one piece, series,
&e. <A simple pistil is of one carpel;
a simple leaf, of one blade, &e.

Simple Fruits, 291.

Simplicissimus. Most simple; complete-
ly simple.

Sinistrérse. Turned or directed to the
left; 51, 140.

Sinuate (-atus). With a strongly wavy
or recessed margin; 98.

Sinus. <A recess or re-entering angle.
Slashed. Same as Laciniate.
Smooth. Either opposed to scabrous,

i. €. not rough, or to glabrous, 7. e.
not pubescent ; the former is the more
correct application.

Soboles. Shoots, especially those from
the ground.

Soboliferous (-us).
lithe shoots.

Solid Bulb. A corm; 61.

Solitary (-arius). Single, only one from
the same place.

Bearing vigorous

Solubilis. Separating into portions or
pieces.
Solitus. Loosed; becoming separate.

Sordidus. Of a dull or dirty hue.

Sorcdiate (-atus). Bearing small patches
on the surface.

Sorema. A heap of carpels belonging to
one flower; 2635.

Sori, sing. sorus. Heaps, such as the
clustered fruit-dots of Ferns.

Sorose. Heaped or bearing Sori.

Sorésis. A fleshy multiple fruit, such
as a mulberry, bread-fruit, and pine-
apple.

Spadiceus. A bright and clear brown,
or chestnut color.

454

Spadiceous.
bearing a

Spadic. A spike with a fleshy axis; 149.

Span. The length of the space between
the tip of the thumb and that of the
little tinger, when outstretched ; about
nine inches.

Sparsus. Sparse or scattered; whence
Sparsiflorus, with scattered flowers;
Sparsifvlius, with scattered leaves, &c.

Spathaceous (-us). Spathe-bearing, or
of the nature of a

Spathe (Spatha). A large bract, or a
pair of bracts, enclosing a flower-clus-
ter; 142.

Spathella. An unused name for the
glumes of Grasses.

Spathilla.
spathe.

Spatulate (Spathulatus). Oblong with
the lower end attenuated, shaped like
a druggist’s spatula; 96.

Species. The particular kind, the unit
in natural history classification; 317.

Specific Character, Name, &c., 349,
363.

Spermaphore or Spermophore (-orum).
A name for the Placenta.

Spermoderm (-ermis). The outer seed-
coat; 305.

Spermodophorum or Sperméphorum. An
unused name for the gynophore in
Umbellifere. The latter also an un-
used name for the Placenta; 261.

Spermothéca. An unused name for peri-
carp.

Spermum. Latin form of the Greek word
for seed. Lat. Semen.

Sphalerocarpium. Name proposed for
an accessory fruit, such as that of
Shepherdia, in which an akene is
enclosed in a baccate calyx-tube.

Spica. See Spike.

Spicate (-atus). In the form of or resem-
bling a spike, or disposed in spikes.

Spiciform (-ormis). Spike-like.

Spicula. .A diminutive or secondary
spike; a Spikelet.

Spike (Spica). A form of indeterminate
inflorescence, with flowers sessile on
an elongated common axis; 149.

Spikelet (Spicula). A secondary spike;
the name given to the Locusta or clus-
ter of one or more flowers of Grasses
subtended by a common pair of
glumes.

Spindle-shaped. See Fusiform.

Spine (Spina). A sharp-pointed woody
or indurated body,commonly a branch,

Having the nature of or

A secondary or diminutive

GLOSSARY.

sometimes a petiole, stipule, or other
part of a leaf; 55, 117.

Spinescent (-ens). Ending in a spine
or sharp point; 59.

Spinose (-osus). Furnished with spines,
or of a spiny character; 55.

Spinuliferous or Spinulose (-osus). Fur-
nished with diminutive spines or Spin-
ule.

Spiral (Spirdlis). As if wound round
an axis. Spiral Ducts, 68. Spiral
Phyllotaxy, 119, 121.

Spiricles. The delicate coiled threads
in the hairs on the surface of certain
seeds and akenes, which uncoil when
wet; 307.

Spithameus. A span long; the length
spanned between the tip of thumb and
forefinger when extended.

Splendens. Resplendent or glittering.

Spongelet, Spongiole (-icla). Name
given to young root-tips; once sup-
posed to be a peculiar organ; 28.

Sporadic (-icus). Widely dispersed or
scattered.

Sporangium. A spore-case or theca con-
taining the analogues of seeds (spores)
in the higher Cryptogams.

Spore (Spora, Greek for seed).
analogue of seed in Cryptogams.

Spore-case. See Sporangium.

Sporidium. Synonym or diminutive of
Spore.

Sporiferous. Spore-bearing.

Spérocarp (-arpium). Name given to
certain spore-cases, as of Lycopodi-
ace.

Sporophore (-orum).
nyms of Placenta.

Sporule (Spdérula). Diminutive spore or
a sort of spore.

Sporuliferous (-us).
taining spores.

Sport. A bud-variation or seed-varia-
tion; 319.

Spumescent (-ens), Spumose.
in appearance.

Spur. <A hollow and slender extension
of some portion of the blossom, usu-
ally nectariferous, as of the calyx of
Larkspur and the corolla of Violet:
rarely applied also to a solid spur-like

The

One of the syno-

Bearing or con-

}Froth-like

process.

Spurred. Producing a spur. See Cal-
carate.

Squama. <A scale of any sort, usually

the homologue of a leaf.
Squamate (-atus), Squamiferous, Squa-
mosus. Furnished with scales. .

aeale

GLOSSARY. 425
Squamella, Squamul. Diminutive compounds as Distichous, two-ranked ;

squama; scales of secondary order
or reduced size.

Squamiform (-ormis).

Squamulose (-osus).
with minute scales.

Squarrose (-osus). Literally rough-
scurfy; applied to bodies rough with
spreading and projecting processes,
such as tips of bracts, Ke.

Squarrulose (-osus). Diminutively squar-
Tose.

Stachys. Greek for spike.

Stalk. Any kind of lengthened support
on which an organ is elevated.

Stamen. One of the elements or phylla
of the andreccium ; 165.

Staumineal, Stamineous (-eus).
to the stamens; 151.

Staminifcrous (-us). Stamen-bearing.

Staminodium. A sterile stamen, or
what answers to a stamen, whatever
its form, without anther.

Staminody. Name for the metamor-
phosis of other floral organs into
stamens; 174.

Standard. The posterior petal of a
papilionaceous corolla; 184.

Seale-like.
Covered or beset

Relating

Stans. Supporting itself in an erect
position.
Station. Particular place as to soil, ex-

posure, &e., which a plant affects;

Stellate (-atus). Star-shaped, arranged
like the rays or points of a star.

Stellulate (-atus) or Stellular. Dimin-
utive of Stellate.

Stem. The main ascending axis of a
plant; 45.

Stemless. See Acaulescent; with no leaf-
bearing stem above ground; 45.

Stemlet. Diminutive stem; as that of
the plumule.

Stenos. Greek for narrow; hence

Stenophyllus. Narrow-leaved, &c.

Sterigma. Any foliaceous prolongation of
the blade of a leaf down on the stem
by decurrence.

Stériymum. Name of Desyaux for the
Dieresilis of Mirbel.

Sterile (-ilis). Barren. as a blossom
destitute of pistil, 191; a stamen
without anther, or an anther without
pollen; an ovary, without good ovules,
seeds without embryo, &e. In com-
mon English use. a male or staminate
‘lower is said to be a sterile flower.

Stichvs. Greek for row or rank, usually
meaning vertical rank ; hence such

Tristichous, three-ranked, &c.

Stigma, pl stigmata. That part or sur-
face of a pistil (usually on or a part of
the style, or in place of it) which re-
ceives the pollen for the fecundation
of the ovules; 166.

Stigmatic (-icus), Stigmatose (-osus).
Relating to stigma.

Stigmatiferous. Stigma-bearing.

Stings. Stinging hairs, seated on a
gland which secretes an acrid liquid,
as in Nettles.

Stipe (Stipes). A stalk of various sorts;
the support of the cap of a mush-
room ; the leafstalk of a Pern ; any
stalk-like support of a gyncecium or
a carpel; 212.

Stipel (Stipelium). An appendage to a
leaflet analogous to the stipule of a
leaf; 106.

Stipellate (-atus). Provided with stipels ;
106.

Stipitate (-atus).
special stalk.
Stipiiifjorm (-ormis). Shaped like a stipe;

stalk-like.

Stipulaceous (-eus), Stipular (-aris).
Belonging to stipules.

Having a stipe or

Stipulate. Possessing stipules.
Stipules. Appendages or adjuncts of a

leaf one ou each side of the insertion ;
85, 105.

Stirps, pl. stirpes. A race.

Stock. Synonym of Race; also the
portion of a stem to which a graft is
applied; a caudex, rhizoma, or root-
like base of a stem from which roots
proceed; 51.

Stole, Stolon (Stolo). A sucker, runner,
or any basal branch which is disposed
to root; 53.

Stoloniferous (-us). Sending of or propa-
gating by stolons, runners, &e.

Stoma, pl. stémata, Stomate. One of
the apertures in the epidermis of folia-
ceous parts, through which cavities
within communicate with the external
air; 89.

Stomatiferous (-us). Bearing stomata
or ‘breathing pores.”

Stone. The hard endocarp of a drupe.

Stone-fruit. A Drupe, such as a peach
or plum: 297.

Stool. The plant from which layers are
propagated, by bending down to the
ground to be rooted.

Stramineous (-eus). Straw-like or straw-
colored.

436

Strap-shaped. See Ligulate (247) and
Lorate.

Striate (-atus). Marked with fine longi-
tudinal lines, streaks, or diminutive
grooves or ridges (Strie).

Strict (Strictus). Close-or narrow and
upright; very straight.

Striyillose (-osus). Minutely strigose.

Strigose (-osus). Beset with strige, or
sharp-pointed and appressed straight
and stiff hairs or bristles.

Strobilaceous (-cus), Strobiliform (-or-
mis). Relating to or 1esembling a
Strobile.

Strobile (Stidbilus). An inflorescence
formed largely of imbricated scales,
as that of Hop and a Fir-cone;
303.

Strombuliformis, Strombuliferus. Twist-
ed spirally into a serew shape, as the
legumes of the Screw-bem (Proso-
pis, sect. Strombocarpa) and of some
species of Medicago.

Strdéphiole (-iola). An appendage ‘at
the hilum of certain seeds; 308.

Structural Botany, 2

Struma. A wen or any cushion-like
swelling on an organ.

Strumose (-osus), Strumiferous (-us).
Furnished with a struma or goitre-like
swelling.

Stupose (-osus). Tow-like; with tufts or
mats of long hairs.

Style (Stylus). The usually attenuated
portion of a pistil or carpel between
the ovary and stigma; 166.

Styliform (-ormis). Style-shaped.

Styliferous. Style-beariug.

Stylinus. Belonging to the style.

Stylosus. With styles of remarkable
length or number, &e.

Stylopodium. Anenlargement ora disk-
like expansion at the base of a style,
as in Umbellifere.

Sub. In composition of Latin words in
terminology, denotes somewhat or
slightly; as, Subacute, Subcordate, that
is acutish, somewhat cordate, Ke.

Subclass, 327.

Subconvolute and Subimbricate in eestiva-
tion, 137.

Suberose (-osus). Of a corky texture.

Subgenus, 327.

Submerged, Submersed (-us).
ufder water.

Suborder (Subordo), 327.

Subpetiolar (-aris). Under the petiole,
as the leaf-buds of Platanus; 42.

Subsection, 327.

Growing

|
|
|
|

}

GLOSSARY.

Subspecies. A group which is ambigu-
ous in rank between variety and spe-
cies; 320.

Subtribe (Subtribus), 327.

Subulate (-atus), Subuliform (-ormis).
Awl-shaped.

Subvariety, 527.

Succise (-isus). As if eut or broken off
at the lower end.

Succubous (-us). When in_ leaves
crowded on a stem the apex of each
leaf is covered by the base of the
next above.

Succulent (Succosus). Juiey.

Sucker. A shoot of subterranean ori-
gin; 53.

Suffrutescent (-ens).
scurely shrubby; 50.

Suffrutez. An undershrub.

Suffruticose (-osus). Low and shrubby
at base; 50.

Suffultus. _Underpropped or supported.

Sulcate (-atus). Grooved or furrowed.

Super. Above. See Supra.

Superior, Superus. Growing or placed
above; also in a lateral flower on the
side next the axis; thus the poste-
rior or upper lip of a corolla is the
superior; 160, 183.

Superposed (Superpositus).
over some other part.

Superposition, 179, 195.

Supervolute (-us), Supervolutive (-ivus).
Same as Convolute when applied to
plaits; 139. 7

Supine (-inus). Lying flat with face up-
ward.

Suppression. Complete abortion ;~179,
190.

Supra. Above; hence in Latin com-
pounds, Supra-axillary, above the
axil; Suprafoliaceous,above a leaf, &e.

Supradedecompound. Several times com-
pound.

Surculose (-osus). Producing suckers.

Surculus. A sucker; a shoot rising from
a subterranean base; 53.

Sursum. Upward; directed upward or
forward.

Suspended (Suspensus). Hanging di-
rectly downward; hanging from the
apex of a cell.

Suspensor of the embrvo, 284.

Sutural (-alis). Relating to a suttire.

Suture (-ura). A junction or seam of
union; used commonly as a line of
opening; 260. e

Sword-shaped. A blade with two sharp
and neaily parallel edges, as in Iris.

Slightly or ob-

Vertically

GLOSSARY.

Syconium or Syconus. A multiple fruit
like that of the Fig; 148, 303. :

Sylvestrts. Growing in woods.

Symmetrical. Regular as to number of
parts or as to shape. In the blossom
it denotes the former; 175.

Symmetry. In the flower relates to
symmetrical disposition of orgaus on
the axis; 174.

Sympetalous (-us). With united petals;
same as Gamopetalous; 244.

Symphiantherous (-us). Same as Sy-
nantherous and.Syngenesious.

Symphysis. Same as Coalescence.

Symphystemonous. With stamens united.

Sympode, Sympodium. A stem made
up of a series of superposed branches
in a way to imitate a simple axis; a
Sympodial stem; 55, 154.

Synacmy. Same as Synanthesis.

Synantherous (-us). Stamens coalescent
by their anthers.

Synanthesis. The simultaneous anthe-
sis or readiness of the anthers and
stigmas of a blossom; 219,

Syncarp, Syncarpium. A multiple fruit
such as a mulberry, or a fleshy aggre-
gate fruit, like that of Magnolia; 299.

Syncarpous (-us). Composed of two or
more united carpels; 261, 263.

Syncotyledonous. With cotyledons sold-
ered together.

Synedral. Growing 6n the angles.

Synema. The column of monadelphous
filaments, as in Mallow.

Syngenesia. Linnzan class (335) charac-
terized by having the anthers united or

Syngenesious. With anthers cohering
in aring; 250.

Synonym. A superseded or unused
name; 354, 365.

Synonymy All that relates to syno-
nyms; 365.

Synsepalous (-us). Of coalescent sepals ;
same as Gamosepalous; 244.

Systematic Botany, 2.

Systylus. The coalescence of styles into
one body.

Tabescent (-ens).
ling.

Tail. Any long and slender terminal
prolongation.

Taper-pointed. See Acuminate.

Tap-root. A primary descending root
forming a direct continuation from the
radicle; 31.

Tawny. Sameas Fulyous; dull brown-
ish-vellow.

Wasting or shrivel-

= Tendril.

|

437

Taxology, Taxonomy. Relating to clas-
sification and its rules; 3, 315.

Teeth. Any small marginal lobes.

Teymen. The inner coat of a seed; 306.

Tela. Latin name for tissue, cellular
tissue, Ke.

Teleianthus. Same as perfect, or her-
maphrodite-flowered.

A filiform production (either
axile or foliar) by which a plant may
climb; 54.

Tepal (Tepalum). <A division of peri-
anth, whether sepal or petal (hardly
ever used).

Teratclogical. Relating to malforma-
tion or monstrous conditions.

Teratology. The science of monsters
and malformations; 170.

Terete (Teres). Round in the sense of
having a circular transverse section.

Tergeminate (-atus). Thrice twin.

Terminal (-alis). Proceeding from or
belonging to the end or apex; 7.

Terminology. Same as Glossology ; 3,
359.

Ternary (-arius). Same as Trimerous ;
consisting of three; 176.

Fernate (Ternus, Ternatus). In threes;
as three in a whorl or cluster.

Tessellated (-atus). In chequer-work.

Testa. The outer seed-coat, which is
commonly hard and brittle, whence
the name, which answers to seed-shell ;
305.

Testaceous (-eus). Of the color of un-

glazed common (brownish-yellow)
pottery.
Tetra. In Greek compounds, four ;

hence
Tetracarpellary (-aris).
pels; 261.

Of four car-

Tetracamarous (-us), Tetracoccus. Of
four closed carpels.
Tetradyndmia. Linnean class (335)

which has the stamens.
Tetradynamous (-us). With four long
and two shorter stamens; 250.

Tetragonal or Tetragunous (-us). Four-
angled.
Tetragynia. Linnean artificial order

(337), characterized by having the
gynecium.
Tetrdgynous. Of four carpels or styles.
Tetramerous (-us). Composed of four
members in a circle; 176.

Tetrandria. Linnean class having the
flowers perfect and

Tetrandrous. With four stamens; 249,
334.

438

Tetrapetalous (-us).
244,

Tetraphyllous (-us).

Tetraquctrous (-us).
or salient angles.

Tetrasepalous (-us).
244.

Tetrdstichous (-us). In
ranks.

Thalamiflorous (-us), 340.
of the flower hypogynous, or on the
Thalamus. The receptacle of a flower;

167. See Torus.

Thallophytes ( Thallophyta), 341.

Thallus. A stratum, in place of stem
and foliage.

Theca. Acase: an anther-cell (251);
a spore-case, &c. (An early name for
the anther, 166.)

Thecaphore (-orum).

With four petals;

Four-leaved; 245.
With four sharp

With four sepals;

four vertical

The stipe of a

earpel (homologous with petiole);
212.
Thorn. Same as spine; 55.

Throat. ‘The orifice of a gamopetalous
corolla or calyx, including any por-
tion between this and the proper tube;
246. See Faux. :

Thyrse, Thyrsus. A contracted or ovate
panicle; a mixed inflorescence, with
inain axis indeterminate, but the sec-
ondary or ultimate clusters cymose ;
15).

Tigelle, Tigellula. A miniature or ini-
tial stem ; sometimes applied to Cauli-
cle (Radicle), sometimes to Plumule;
10.

Tinctorius. Dyed; used for dyeing;
imparting color.

Tissue. The anatomical fabric.

Tomeéntose (-osus). Densely pubescent
with matted wool, or Tomentum.

Tongue-shaped. Long and nearly flat,
somewhat fleshy, and rounded at the
apex.

Tooth. See Teeth.

Tvothed. See Deutate.

Top-shaped. Junversely conical.

Torose (-osus). Cylindrical, with con-
tractions or bulges at intervals.

Tortuous (-osus). Bent or twisted in
different directions.

Torulose (-osus).
slightly torose.

Diminutively or

Tortus. Twisted.

Tortilis. Susceptible of twisting.

Torus. The receptacle of a flower; 167,
211.

Trabeculate (-atus). Cross-barred.
Trachea. Aspiral vessel or duct, named

With parts

GLOSSARY.

from resemblance to the trachee of
insects.

Trackycarpous (-us) Rough-fruited.

Trachyspermous. Rough-seeded, &e.

Transverse (-ersus). Across; right and
left as to bract and axis; collateral;
160.

Trapeziform (-ormis), Trapezoid. Un-
symmetrically four-sided, like a tra-
pezium.

Tree. A woody plant with an elevated
trunk.

Tri-. In compound words, both Latin
and Greek, denotes three or triple.

Triachanium. A fruit like a cremocarp
but of three carpels.

Triadelphous, Triadelphia.
iments in three sets; 250.

Triandria. Linnean class (3384) with the
flowers.

Triandrous. With three stamens; 249.

Triangular (-aris), Triangulatus. Three-
angled.

Trianthous (-us). Three-flowered.

Tribe. Group superior to genus, in+
ferior to order; 326.

Tricarpellary (-aris).
261.

Tricarpous (-us).
fruits or carpels.

Tricephalous (-us). Bearing three heads.

Trichocarpous (-us). Hairy-fruited.

Trichodes. Resembling hair.

Trichdtomous (-us). Three-forked ;
branched into three divisions.

Tiichome (Trichéma). Any outgrowth
of the epidermis, such as a hair or
bristle; 209.

Tricoccous (-us).
cocci.

Tricolor. Three-colored.

Tricuspidate (-atus). Tipped with three
cusps or pointed tips.

Tridéntate (-atus). Three-toothed.

Tridigitate. Thrice digitate.

Triduus. Lasting for three days.

Tricnnial (Triennis). Lasting for three
years.

Tiifarious (-ius). Facing three ways,
in three vertical ranks.

Trifid (Ti ifidus). Three-cleft.

Trifoliate (-ius, Trifoliatus).
leaved.

Trifoliolate (-atus).

Trifiircate (-atus).
forks or branches.

Trigamous (-us). Bearing three kinds
of flowers.

Tiigonous (-us), Trigonal. Three-angled.

With fila-

Of three carpels;

Consisting of three

Consisting of three

Three-

Of three leaflets.
Divided into three

GLOSSARY.

Trigynia. Linnean artificial order with
Trigynous, zi. e. three-styled flowers;
337.

Trihilatus. Waving three apertures, as
in some grains of pollen.

Trijugate (Trijugus). With three pairs
of leaflets or pinuz.

Trilobate (Trilubus). Three-lobed.

Trilocular (-aris). Three-celled.

Trimerous (-us). Three-membered parts
in threes; 176.

Trimestris. Lasting for or maturing in
three months.

Trimorphous, Trimorpism.
under three forms; 236.

Trinervate (Trinervius).

Trinodal. Of three nodes or joints.

Triecia. Linnean artificial order with
the flowers.

Triecious or Trivicous (-us). Having
staminate, pistillate and perfect flowers
(or three kinds of flowers as to sex),
334; on three distinct plants.

Tridvulate (-atus). Having three ovules.

Tripartible (-ibilis). Tending to split
into three portions.

Tripartite (-itus). ‘Three-parted.

Tripetalvid (-videus). As if three-pet-
alled.

Tv ipetulous(-us). Having three petals.

Triphyllous (-us). Three-leaved; 243.

Tripinnate (-atus). Thrice pinnate; 104.

Tripinnatifid (-idus). Thrice pinnatifid.

Triple-ribbed or nerved. With midrib
dividing into three, or sending off on
each side a strong branch, above the
base of the blade; 93.

Triplinerved (Triplinervius). Same as
Triple-nerved, Triple-ribbed; 93.

Tripterous (-us). Three-winged.

Triquetrous (Triqueter). Three-edged ;
with three salient angles.

Triquinate (-atus}. Divided first into
three then into five.

Trisected (-us). Divided into three por-
tions; 99.

Trizepalous (-us). Of three sepals.

Triserial (-alis), Triseriate (-atus). In
three horizontal ranks or series.

Tristachyus. Three-spiked.

Tristichous (-us). In three vertical
ranks; 122.

Tristigmatic. With three stigmas.

Tristis. Dull colored.

Tristylous (-us). Having three styles.

Trisilcate (-atus). Three-grooved.

Triternate (atus). Thrice ternate; 104.

Trivial names, Nomina trivialia. Com-
mon or yulgar names; used by Lin-

Occurring

Three-nerved.

439

neus for specific names of a single
word; 346, 362.

Trochlear (-earis). Pulley-shaped.

Trophosperm (Trophospermium). Name
for the Placenta; 261.

Trumpet-shaped. ‘Tubular, with a dilat-
ed orifice.

Truncate (-atus).
end; 97.

Trunk (Truncus). A main stem.

Tryma. A drupaceous nut, with exo-
carp at length dehiscent or otherwise
separating, such as walnut and hick-
ory nut.

Tubeformis. Trumpet-shaped.

Tube (Tubus). Any hollow elongated
body or part of an organ; 245.

Tuber. A thickened and short subter-
ranean branch, beset with buds or
eyes; 59.

Tubercle (Tuberculum). A small tuber
or an excrescence: or something be-
tween a tuber and a root; 60.

Tuberculate (-atus). Beset with knobby
projections or excrescences.

Tuberiferous. Bearing tubers.

Tubular, Tubulosus (-ose). Having a
tube; tube-shaped; 248.

Tubuliflorus (-us). When the flowers of
a head have only tubular corollas.

Tunicate (-us). Having coats (tunics).

Turbinate (-atus). Top-shaped.

Turion, (Turio). A scaly sucker or
shoot from the ground; 41.

Turnip-shaped. See Napiform.

As if cut off at the

Twin. In pairs. See Geminate, Didy-
mous.
Twining. Winding -spirally and so

climbing (Twiners); 51.
Twisted. Contorted.
Two-lipped. See Bilabiate.
Type. The ideal plan or pattern.
Typical. Representing the plan or
type.

Uliginose (-osus). Growing in swamps.

Ulnaris. Of the length of the ulna or
fore-arm.

Umbel (Umbélla). An inflorescence
(properly of the indeterminate type)
in which a cluster of pedicels spring
all from the same point, like rays of
an umbrella; 146.

Umbellate (-atus). Umbellitorm (-ormis).
In or like umbels.

Umbellet. A partial or secondary um-
bel; 150.

Umbellifercus (-us). Bearing umbels.

440 GLOSSARY.

Umbéllula. A partial or secondary um-
bel, or umbellet; 150.

Umbilicate (-atus). Depressed in the
centre, navel like.

Umbilicus. The hilum of a seed.

Umbonate (-atus). Bearing an Umbo
or loss in the centre

Umbraculiform (-ormis). Having the
general form of an umbrella. ,

Umbrosus. Growing in shady places.

Unarmed. Destitute of prickles, spines,
or other armature.

Uncate (-atus), Uncinate, (-atus), Unei-
form (-ormis). Hooked; bent or
curved at tip in the form of a hook.

Uncialis. An inch (uncia) in length.

Undute (-atus) or Undulate (-tus).
Wavy; 98.

Undershrub. A very low shrub; 50.

Unequally pinnate. See Impari-pin-
nate.

Unguiculate (-atus). Contracted at
base into an

Unguis. A claw, or stalk-like base of a
petal, &c.; 245.

Uni-. In Latin compound, one; as

Unicellular. Of one cell; Unicolor, of
one color, &e.

Unicus. Singly or single, solitary.

Uniflorous (-us). One-flowered.

Onifoliate (-atus). One-leaved.

Unifoliclate, of one leaflet; 102.

Unijigate (Unijugus). Of one pair;
102.

Unilabiate (-atus}. One-lipped, like the
corolla of Acanthus, in which the
upper lip is obsolete.

Unilateral (-alis). One-sided; either
originating on or more commonly
turned all to one side of an axis.

Onilocular {-aris). One-celled.

Uninervate (Uninervis, Uninervius).
One-nerved.

Uniovulate (-atus). Having only a soli-
tary ovule. =

Uniparous. Bearing one; as a cyme of
one axis or branch; 152, 155.

Uniserial (-ialis), Uniseriate (-atus).
In one horizontal row or series.

Unisexual (-alis, Uniserus). Of one
sex; haying stamens only or pistils
only; 191.

Univalved ( Univalvis). Of one piece or
valve.

Ureevlate (-atus) Hollow and con-
tracted at or below the mouth, like an
urn or pitcher ( Urceolus).

Urens. Stinging, in the manner of net-
tles.

Utricle (Utriculus}. A small bladdery
pericarp; 295. Orany small bladder-
shaped body or appendage; also a
synonym of a cell of parenchyma.

Utricular (-aris), Utrieulate (-atus),
Utriculiform (-ormis), Utriculose
(-osus). Havirg or consisting of
utricles, or bladder-like in appear-
ance.

Vacillans. Swinging free, as the anth-
ers of Grasses on their filaments.

Vacuus. Void or empty of the proper
contents.

Vagina. A sheath, as of a leaf, &e.

Vaginate. Sheathed.

Vall cule. The intervals or grooves
between the ridges or ribs of the fruit
Umnbellifere.

Valcate (-atus), Valvular (-aris).
Opening as if by doors or valves, as

do most dehiscent fruits (capsules),

and some authers; also the parts of
a flower-bud when they exactly meet
without overlapping; 135.

Valve (Valva). One of the pieces into
which a capsule splits, 288.

Valved. Same as valvate: hence 3
-valved, 5-valyed, many valved, &e.
Valeula. A diminutive valve. Also
used (after Linnzus) for the inner or

flower-glumes of Grasses.

Variegated (-atus). Irregularly colored ;
in patches of color.

Variety (Varietas). A sort or modifi-
cation subordinate to species; 318.

Varivlate, Variolaris, Marked as if
by the pustules or pittings of small-
pox.

Vascular (-aris). Relating to or fur-
nished with vessels ( Vasa) or ducts.

Vascular Plants ( Vasculares), 340.

Vasculum. Same as Ascidium. Also
the botanists’ collecting box ; 372.

Vasiform (-ormis). In the form of a
vessel, duct, &e.

Veined. Furnished or traversed with
fibro-vascular bundles or threads, es-
pecially with those which divide and
are reticulated.

Veins (Vene). In general any ramifi-
cations or threads of fibro-vascular
tissue in a leaf or any flat organ;
especially (as distinguished from
nerves) those which divide or branch ;
92.

Veinless. Destitute of veins.

“einlet (Venula). Gne of the ultimate

ow fe — wren tet pn” * *

1 cnt er INN aoa =

on

GLOSSARY.

or smaller ramifications of a vein or
rib; 93.

Velute (-atus). Veiled.

Veliutinous (Velutinus). Velvety: the
surface covered with a soft coating of
fine and close silky pubescence, or
velumen.

_ Venation ( Venatio).
ing; 90.

Venenatus, Venenosus. Poisonous.

Venose (-osus) Veiny; abounding in
veins or network.

Ventral (-alis). Belonging to the an-
terior or inner face of a carpel, Kc.;
the opposite of dorsal.

Ventricose (-osus). Swelling unequally
or inflated on one side.

Ventriculose (-osus). Minutely ventri-
cose. :

Venulose (-osus). Abounding with vein-
lets or venule.

Vermicular (-aris) | Worm-shaped.

Vernal (Vernalis), Appearing in spring.

Vernation (-ativ). The disposition of
parts in a leaf-bud; 132.

Vernicose (-vosus). As if varnished.

Veérrucose (-osus). Covered with warts
(verruce) or wart-like elevations.

Versatile (Versatilis). Swinging toand
fro; turning freely on its support;
253.

Versicolor. Changing color, or of more
than one tint cr color.

Vertex. The apex of au organ.

Vertical (-alis). Perpendicular to the
horizon; longitudinal.

Verticil (-illus). A whorl; 6.

Verticillaster. A false whorl, composed
of a pair of opposite cymes; 159.

Verticillastrate. Bearing or arranged
in Verticillasters.

Verticillate (-atus, -aris).
whorl; 6, 119, 120.

Vescicle (-icula). A small bladder or
air-cavity.

Vesicular (-aris), Vesiculose (-osus). As
if composed of little bladders.

Vespertine (Vespertinus). Appearing or
expanding in early evening.

Vessels (Vase). See Ducts.

Verillary (-aris), Vesxillar, 137. Per-
taining to the

Vezillum. The standard or large pos-
terior petal of a papilionaceous corolla;
184. ©

Villose (osus) or Villous. Bearing shaggy
or long and soft (not interwoven) hairs
or Villi.

Vimineous(-eus). Bearing long and flex-

The mode of vein-

Disposed in a

441

ible twigs, like those used for wicker
work. :

Vine. Any trailing or climbing stem:
originally that of the Grape from
which wine is made.

Vinealis. Growing in vineyards.

Violaceous (-eus). Violet-colored.

Virens. Green, or evergreen.

Virescens. Greenish or turning green.

Virgate (-atus). Wand-shaped, or like
a rod; slender, straight, and erect.

Virgultum. A vigorous twig or shoot.

Viridescent (-ens). Same as Virescens.

Viridis. Green.

Viridulus. Greenish.

Virosus. Yenomous.

Viscid (-idus), Viscous (-osus). Sticky
from a tenacious coating or secretion.

Vitellinus. The yellow hue of the yolk
of egg.
Vitéllus. Name formerly given to the

peculiar albumen which is in some
cases deposited within the embryo-sac.

Viticulose (-osus). Sarmentaceous ; pro-
ducing vine-like twigs or suckers,
viticule.

Vitte. The fillets or stripes (vil-tubes)
of the pericarp of most Umbelliferz,
which contain an aromatic or peculiar
secretion.

Vittate (-atus). Bearing vittee; or with
any longitudinal stripes.

Viviparous (-us). Germinating or sprout-
ing from seed or bud while on the
parent plant.

Volutle ( Volibilis).
support; 51.

Volutus. Rolled wp in any way.

Volva. A wrapper or external covering,
especially that of many Fungi.

Twining round a

Wavy. See Undulate.

Waszy. Resembling beeswax in consist-
ence or appearance.

Wedge-shaped or Wedge-form. See
Cuneate; 95.

Wheel-shaped. See Rotate.

Whorl. Arranged in a circle round an
axis; a Verticil; 6.

Whorled. Disposed in whorls.

Wild. Growing without cultivation;
spontaneous.
Winy. See Ala. Any Membraneous

or thin expansion by which an organ
is bordered, surrounded, or otherwise
augmented. Also the two lateral
petals of a papilionaceous corolla are
termed wings; 185.

442

Winged. See Alate: bearing a wing or
wings.

Withering. See Marcescent.

Wood. The hard part of a stem, &c.,
mainly composed of

Wovod-cells, Woody fibre or tissue, 68.

Woolly. See Lanate and Tomentose:
clothed with long and tortuous or
matted hairs.

Xanthos. Greek for yellow in com-
pounds, such as Xanthephyll, the yel-
low coloring matter in leaves.

GLOSSARY.

Xenogamy. Fecundation of the ovules of
a flower by pollen from some other
plant of the same species; cross-fer-
tilization; 216.

Xylinus. Woody, pertaining to wood.

Zoospore. One of the free-moving spores
of the lower Cryptogams.

Zygomorphous (-us). That which can be
bisected in only oue plane into similar
halves; 175.

ADDENDA.

Antidromous, Antidromy.
Infertile (-iiis).

When the course of a spiral is reversed, 157.
Said of a pistil or flower which fails to set fruit.

Polyembryony. The production of two or more embryos in a seed, 284.

Saprophytes (-yta).

Plants feeding upon decaying vegetable or animal matter.

J
r <5
. 7
¥ ey
ye ots ¥
~ - | Cae bw o
Soe . ; » 4 ¢ ;
ee put jhe Mates Aa
a > |
> in - =
iy Z Sy Ae
é * = *
pa a fee tH fa , oe. oan 7
es ¢ *
<a S ny = —
: alt a, ‘ ps
_— %, .
re 2s mS I Sole *
er
>.
4 eb ~~.
Pa .
a - +
-_
.
~
=
«
-
= a
“ “a
s
’
-» 7 « .
‘
~ .
-
.

.
f
~
_*
]

UNIVERSITY OF TORONTO
LIBRARY

—_—

Do not

re move

the card |

\
\

from this \_

Pocket.

———_

Acme Library Card Pocket
Under Pat. ‘‘ Ref. Index File.”
a Made by LIBRARY BUREAU

0 600 OL 980 SGI 60 6€
2 WALI SOd 4SIHS AVE JONVY

MAIASNMOG LV